oviposition selection by a rare grass skipper, polites mardon, in

Transcription

OVIPOSITION SELECTION BY A RARE GRASS SKIPPER, POLITES MARDON, IN
MONTANE HABITATS: ADVANCING ECOLOGICAL UNDERSTANDING FOR
DEVELOPING CONSERVATION STRATEGIES
By
LONI JEAN BEYER
A thesis submitted in partial fulfillment of the requirements for the degree of
MASTER OF SCIENCE IN ENVIRONMENTAL SCIENCE
WASHINGTON STATE UNIVERSITY VANCOUVER
School of Earth and Environmental Science
MAY 2009
To the Faculty of Washington State University:
The members of the Committee appointed to examine the thesis of
LONI JEAN BEYER find it satisfactory and recommend that it be accepted.
__________________________________
Chair
__________________________________
__________________________________
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ACKNOWLEDGEMENTS
In addition to my committee members Cheryl Schultz, Scott Black and Brian
Tissot, I would like to thank my fellow lab mates Aldina Franco, Dina Roberts, Caitlin
LaBar, Erica Henry, Leslie Rossmell, Cheryl Russel, Alexa Carelton for constant
feedback and support. A special thanks to John Scott, Barbara Wilson, and Wayne
Rolle who were integral to the identification of over sixty graminoid species. I am
grateful for the dedication and diligence of field assistants Luke Frishkoff, Mike
Lumio, and Betsy Scott as our team succeeded in covering expansive territory and
collecting a fantastic amount of data. I am thankful for the support of Tom Kogut,
Mitch Wainwright, Dave Nunnalee, and Ann Potter for sharing their knowledge of the
Gifford Pinchot National Forest and expertise on Polites mardon. Finally, I would like
to thank Kelly Van Norman, Marianne Turley, Wayne Rolle, Jim Algeria, Norm
Barrett, Bruce Marcot, Darci Rivers-Pankratz, and Dana Ross, for their feedback on
protocols and statistics; and the support of the Xerces Society for Invertebrate
Conservation.
This project was funded by the United States Forest Service, the United States
Bureau of Land Management, Washington State University Vancouver, and a WSU
Robert Lane Fellowship in Environmental Studies to Loni J. Beyer.
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OVIPOSITION SELECTION BY A RARE GRASS SKIPPER, POLITES MARDON,
IN MONTANE HABITATS: ADVANCING ECOLOGICAL UNDERSTANDING FOR
DEVELOPING CONSERVATION STRATEGIES
Abstract
by Loni Jean Beyer
Washington State University Vancouver
May 2009
Chair: Cheryl B. Schultz
The Grass skipper subfamily (Hesperiidae) includes many at risk species
across the globe. Conservation efforts for these skippers are hindered by
insufficient information about their basic biology. The rare Washington State
endangered Mardon skipper (Polites mardon) is declining throughout its range. We
surveyed Mardon oviposition across 9 study meadows in the Gifford Pinchot
National Forest of Washington State. We conducted habitat surveys with respect
to oviposition (n=269) and random (n=270) locations, recording data on over 50
variables. Mardon oviposited on 23 different graminoid species, yet are selective
for specific graminoids within meadows. Most frequent ovipositions across
meadows occurred on Festuca idahoensis and Poa pratensis (accounting for 112
of 269 total oviposition observations). Discriminant Function Analyses revealed
that Mardon habitat was too variable to detect oviposition selection across study
meadows, yet there was strong selection occurring within meadows (r2 ranging
from 0.82-0.99). Variables important to within meadow selection were graminoid
cover, height, and community; oviposition plant structure (leaf density, height,
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area); insolation factors (tree abundance and canopy shading); and litter layer
factors (cover and depth). With few exceptions the primary discriminating variables
were significantly different (p<0.001). Conservation implications include
maintaining native meadow ecosystems with sensitivity to local habitat
preferences.
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TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS .......................................................................................iii
ABSTRACT .............................................................................................................iv
LIST OF TABLES....................................................................................................vii
LIST OF FIGURES..................................................................................................viii
INTRODUCTION.....................................................................................................1
METHODS ..............................................................................................................5
STUDY SPECIES & HABITAT .....................................................................5
MEADOWS ..................................................................................................6
OVIPOSITION SURVEYS ............................................................................7
POPULATION COUNTS ..............................................................................8
HABITAT SURVEYS ....................................................................................9
DATA ANALYSIS .........................................................................................11
RESULTS ...............................................................................................................13
DISCUSSION..........................................................................................................16
CONSERVATION IMPLICATIONS ..............................................................20
REFERENCES........................................................................................................24
APPENDICIES ........................................................................................................39
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LIST OF TABLES
Page
TABLE 1: MEADOW DESCRIPTIONS ...................................................................30
TABLE 2: OVIPOSITION PLANT SPECIES ...........................................................31
TABLE 3: OVIPOSITION FREQUENCY BY PLANT SPECIES ..............................32
TABLE 4: WITHIN MEADOW PRIMARY DESCRIPTORS ....................................33
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LIST OF FIGURES
Page
FIGURE 1: RESEARCH MEADOWS......................................................................34
FIGURE 2: QUADRAT AND SUB-PLOT DIAGRAM ...............................................35
FIGURE 3: AMONG MEADOW DFA ......................................................................36
FIGURE 4: WITHIN MEADOW DFA ......................................................................37
FIGURE 5: ACROSS MEADOW TRENDS .............................................................38
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Dedication
This thesis is dedicated to the Mardon skippers (Polites mardon), all those who
have worked toward Mardon skipper conservation, and the friends who have
supported them during this endeavor. Together we can accomplish great things.
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1. Introduction
Lepidoptera are one of the largest, most diverse, and most endangered
taxonomic groups (Liu et al. 2006; Smallidge & Leopold 1997; Thomas et al.
1994). Habitat loss and degradation has led to declines in butterfly populations
across many parts of the world; including Europe, Japan, Asia, Australia, and
North America (Albanese et al. 2007; Bergman 1999; Eastwood & Hughes 2003;
Fox et al. 2006; Freese et al. 2006; Liu et al. 2006; Smallidge & Leopold 1997;
Thomas et al. 1994). Successful recovery of at risk species largely depends on a
sufficient understanding of their basic biology, yet this knowledge is often lacking
for rare butterflies (Schultz & Crone 2008).
Butterfly declines often signal the degradation of the habitats with which
they are associated (Oostermeijer & van Swaay 1998). Lepidoptera have a
polymorphic life history, including a larval form in the juvenile state and a winged
form in the adult state, making them dependent on a variety of resources within
their environment. Adult life stages require sufficient food resources, most
commonly nectar flowers, access to host plants, and large scale structural
components; such as habitat connectivity, refuge from adverse weather, or
adequate insolation (Dennis et al. 2006). Larval stages may require specific plant
species for forage as well as particular micro-habitat conditions (Albanese et al.
2007; Awmack & Leather 2002; Grundel et al. 1998). The dependence on so many
habitat variables creates sensitivity to even small changes within the ecosystem,
and many species are considered environmental indicators (Brown & Freitas 2000;
Eastwood & Hughes 2003; Oostermeijer & van Swaay 1998). Rare butterflies are
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especially useful for monitoring unique ecosystems and are often associated with
other threatened fauna (Brown & Freitas 2000).
An understanding of what factors determine essential habitat for rare
butterflies is imperative to their conservation. Important habitat characteristics are
commonly determined by investigating larval habitat use (Anthes et al. 2008; Ellis
2003). The susceptibility of butterflies to environmental changes is pronounced in
the larval state due to their limited mobility and restricted habitat requirements
(Anthes et al. 2003; Thomas et al. 2001). Larval survivorship is largely determined
by ovipositing females, as larvae generally do not travel far, if at all, from their
natal locations (Awmack & Leather 2002; Bergman et al. 2004; Doak et al. 2006).
Female butterflies are selective during oviposition, depositing eggs in
locations that are favorable to larval development and survival will increase their
fecundity (Awmack & Leather 2002). Correspondingly, female butterflies may
increase the number of eggs deposited in high quality habitats and host plants
(Chen et al. 2004; Fownes & Roland 2002; Mizumoto & Nakasuji 2007). Habitat
factors that a female butterfly may cue in on when selecting a suitable oviposition
location include host plant species (e.g. Mountain Apollo butterfly, Parnassius
Apollo, Fred et al. 2006), the host plant nutritional and chemical content (e.g.
Cabbage white butterfly, Pieris rapae, Chen et al. 2004), the host plant size and
structure (e.g. Marsh fritillary, Euphydryas aurina, Anthes et al. 2003), or
oviposition-location microclimate (e.g. Karner blue butterfly, Lycaeides melissa
samuelis, Grundel & Pavlovic 2007; Grundel et al. 1998).
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The skipper butterfly family, Hesperiidae, harbors approximately 4000
species (Warren et al. 2008). There are at least 55 at risk skippers world wide,
including a minimum of 35 grass skippers (subfamily Hesperiinae, Appendix A).
The Mardon skipper (Polites mardon, U.S. federal candidate, Washington State
endangered) is a rare and declining butterfly in the Pacific Northwest of the United
States of America. The basic biology of this species is poorly understood (Beyer &
Black 2006; Black & Vaughan 2005; Potter et al. 2002). In the U.S. there are three
federally listed skippers, including the Carson wandering (Pseudocopaeodes
eunus obscurus, endangered) Laguna Mountains (Pyrgus ruralis lagunae,
endangered), and Pawnee Montane (Hesperia leonardus montana, threatened).
There are two U.S. federal candidate species the Dakota skipper (Hesperia
dacotae, Canada endangered) and the Mardon skipper (Polites mardon,
Washington State endangered), as well as several other state-listed skippers. To
date, limited information on the habitat requirements inhibits management efforts
for all of these butterflies (Beyer & Black 2006; Potter et al. 2002; USFWS 1997,
1998, 2005; Warren et al. 2008). Grass-feeding butterflies, in general, have highly
complex resource requirements and very little is known about how they utilize their
habitats (Dennis et al. 2006).
Mardon skippers occur across a range of habitat types which vary from
expansive 5,000 ha of low elevation prairies to isolated high elevation forested
meadows that ranging in size from 0.5 to 5 ha (Beyer & Black 2006; Potter et al.
2002). However, the aspects of these habitats critical to Mardon persistence are
entirely unknown. In 2006 we conducted an exploratory study with the Xerces
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Society for Invertebrate Conservation. In this study we investigated oviposition
behavior of the Mardon skipper. Eleven species of grasses and sedges were
observed as oviposition plants (Beyer & Black 2006; Potter et al. 2002). What was
once considered the primary larval host plant, Festuca species, was absent in
many of the known sites. This result completely changed former perceptions about
Mardon habitat (Black & Vaughan 2005), and further stimulated inquiry as to what
makes this butterfly rare.
In this study we investigate Mardon skipper site utilization across a variety
of montane habitat types to determine what aspects are critical to population
recovery. The primary goal of this study is to determine what influences Mardon
skipper oviposition location selection, thereby understanding larval habitat
requirements. We aim to determine (1) what graminoid species are utilized for
oviposition, (2) what landscape and local factors influence oviposition selection,
and (3) to what extent these factors vary between sites. The answers to these
questions are essential to understanding Mardon habitat requirements. This
information is the first step in developing a Mardon skipper conservation plan and
serves as baseline ecological information from which future research can be
founded. Additionally, the information herein contributes to the conservation of
other rare and endangered skippers by advancing knowledge of the understudied
grass skipper family.
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2. Methods
2.1 Study Species & Habitat
The Mardon skipper belongs to the grass skipper subfamily, Hesperiinae.
Skippers of this family forage exclusively on graminoids as larvae, and are
dependent on meadow-grassland habitats. Female Mardon deposit eggs singly
while perched. Distributions of extant Mardon skipper populations are disjunct;
ranging from the grasslands of northwest California to the Puget Tough including
the Cascade Mountain Range in both Oregon and Washington State. All known
Mardon skipper sites are small; most support populations of less than 50
individuals and are isolated from neighboring populations (Black & Vaughan 2005;
Potter et al. 2002).
Existing Mardon habitat has undergone major reductions and recently
several populations have been extirpated (Black & Vaughan 2005). Threats to its
existing habitat are a consequence of urban development, contemporary resource
management practices (logging, grazing, and fire suppression), and increased
recreational use of national forests and public lands (Black & Vaughan 2005).
Montane meadow habitats have drastically declined (Coop & Givnish 2007;
Roland & Matter 2007). Fire suppression over the last century has lead to tree
and shrub encroachment in forest meadows (Norman & Taylor 2005). Grazing,
recreation, and increased logging roads, and agriculture have aided the spread of
invasive weeds (Leung & Marion 2000; Trombulak & Frissell 2000). As a result,
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meadows and grasslands are disappearing (Griffiths et al. 2005) or undergoing
drastic habitat changes (Crawford & Hall 1997; Noss et al. 1995).
2.2 Meadows
Since 2000, thirty-nine Mardon skipper meadows have been documented in
the Gifford Pinchot National Forest. These meadows range from 800 to 1700
meters in elevation, and have various management histories. This provided a
great opportunity to study several distinct populations, each persisting within
different habitat types, and allowing us to determine the commonality between
them. Twenty-one of the 39 documented Mardon skipper meadows had historically
recorded populations where counts exceeded 10 butterflies, and were scouted as
potential research sites. Of these 21 meadows, three were excluded because they
were not logistically feasible, five of the populations never produced adults during
our research season, and four were excluded due to small population sizes (under
15 individuals) during the 2007 season. The nine remaining meadows were
included in this study (Figure 1): Cave Creek, Peterson, Lost, Flog Salvage,
Midway, Smith Butte, Muddy, 7A, and Grapefern.
Dominant vegetation at all study meadows consisted of a mix of grasses
and sedges. Rushes were only noted present at Cave Creek, Muddy, and 7A
meadows which are a mix of moist wetland and dry grassland. Cave Creek is
particularly impacted by noxious weeds; various strategies to control
houndstounge (Cynoglossum officinale) and Canada thistle (Circium arvense)
have been implemented since 2004, including weed-whacking and hand removal.
Approximately 80% of the Cave Creek meadow has been fenced to reduce cattle
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grazing impacts and the spread of invasive weeds. Historically, Peterson Prairie
was fenced to contain livestock, and grazing impacts were heavy. In recent years
all grazing and storing of livestock on Peterson Prairie has ceased. Flog Salvage
was heavily logged historically, and reseeded with Lodgepole pine (Pinus
contorta). Approximately 80% of the original meadow is now densely overgrown
with saplings, and graminoid diversity there includes only a few species. Lost
Meadow, bordered by an open second generation forest, is exposed to short
periods of heavy grazing. Midway is an open meadow connected to other open
areas of potential Mardon habitat. The area surveyed was chosen based on a
priori knowledge of high Mardon use areas.
Microhabitat temperature and humidity affect larval survivorship, and may
influence oviposition behaviors (Kuhrt et al. 2006; Schweiger et al. 2006). We
distributed four to five iButton data-loggers, model DS1923-F5, throughout six of
the study meadows to capture ground level temperature and relative humidity.
Loggers recorded data every four hours for one year (Appendix G).
2.3 Oviposition Surveys
Within each meadow we surveyed Mardon skipper oviposition selectivity.
Surveys were conducted on calm (<5 on Beauford wind scale), sunny days with
temperatures above 15° Celsius. Oviposition observations began when any
individual female butterflies were observed flying. A random point within the
meadow was located, using random number tables, from which a transect line was
walked until a female was encountered. Observations were made with the aid of
8x42 binoculars. If the female being observed was not indicating oviposition
7
behavior after 10 minutes the surveyor terminated the observation. If oviposition
was still suspected to occur then the observation continued for an additional 10
minutes. If no oviposition had occurred at the end of 20 minutes the observation
was terminated, and another female was located from a new random transect.
Females engaged in oviposition were watched for up to five individual egg laying
behaviors. Often the female was lost after exhibiting oviposition as the observer
prioritized marking the oviposition location over continuing observation on the
individual.
All precise physical locations where oviposition occurred, hereafter referred
to as “oviposition locations”, were marked with metal stakes. The number of days
spent surveying oviposition behavior at a single meadow ranged from one to six
days, occasionally spanning a few weeks. We targeted marking a minimum of 30
oviposition locations per study meadow. The duration of time surveying each
meadow was subject to how long it took to meet that target, which was highly
dependant on weather conditions and population size. Thirty random-haphazard
locations were also selected from each meadow. We would determine a starting
point and pacing distance by use of random number tables. Pacing direction would
be determined by indiscriminately selecting a compass bearing. Random “host
plants” were determined by blindly throwing a pin flag from the random-haphazard
locations, hereafter called “random locations”. All variables were recorded in the
same way for both random and oviposition locations.
2.4 Population Counts
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Population counts were conducted every 5-7 days, between 10 AM and 5
PM on sunny days with low wind speeds (<5 on Beauford wind scale) and
temperatures above 15° Celsius. We walked transect lines spaced approximately
10 meters apart. When Mardon were observed the number of butterflies was
recorded, and their spatial locations were recorded on a map or with GPS.
Butterflies that entered from behind the surveyor were ignored. In meadows where
there was overlap in the flight periods of Mardon and closely related Sonora
skippers (Polites sonora), an individual of both species was caught and viewed to
acclimate the observers eye. Thereafter, Mardon skipper identification was made
without capture and with the aid of 8x42 binoculars.
2.5 Habitat Surveys
To capture the environmental conditions at the time of oviposition all habitat
surveys were conducted within 7 days of the observed oviposition for oviposition
locations, and during the meadow-specific adult flight period for the random
locations. Both local and landscape variables influence butterfly oviposition
behavior and larval survivorship (Davis et al. 2007; Kuussaari et al. 2007;
Schweiger et al. 2006). We measured over 50 variables at each random and each
oviposition location to capture as many possible factors that may contribute to
Mardon reproductive ecology (Appendix B). These variables included
characterization of graminoid communities, the oviposition plant, the fine scale
microhabitat, and the meadow landscape.
A one m2 quadrat, centered at each oviposition and random location, was
utilized to capture the local habitat. Each graminoid species was recorded along
9
with its corresponding percent cover and maximum height. Litter depth, soil pH
and soil moisture potential were measured as near to the oviposition location as
possible without disturbing the egg. Soil variables were measured with a Kelway
Soil Tester Model HB-2. Total percent cover of vascular plants, forbs, graminoids,
litter, rocks, shrubs, trees, bare ground, and cryptograms was estimated. The
tallest plant was measured and identified to species. Horizontal vegetation
thickness was measured by recording the percent cover, on a meter stick position
parallel with the ground, at 0.3, 0.6, and 0.9 meter heights.
A 0.1 m2 sub-plot was used to characterize the vegetation community in the
immediate vicinity of oviposition or random locations (Figure 2). Percent cover of
total graminoid and total forbs, as well as the percent cover of each species of
graminoid and their corresponding maximum heights were measured within the
subplot. Graminoid species richness was included as a variable within both the
quadrat and subplot, as well as graminoid heterogeneity and evenness indices
(abundance by percent cover).
Oviposition plants and random “host plants” were identified to species.
Graminoids often grow in bunches of several leaf blades, and occasionally more
than one species is present in those bunches. If it was not clear which species
was being selected by the female skipper, then the other species present in the
bunch were recorded and noted as “mixed”. Oviposition plant (or bunch) length,
width, maximum basal leaf height, and maximum culm height were measured. The
plant density was categorized from 1 to 4 as a solitary blade, a loose structure
(approximately 2-50 blades), a dense structure (50+ blades), or matted (forming a
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carpet like coverage) respectively. The percent of the oviposition plant that was
dead (brown) was estimated. Finally, the distance to the nearest neighboring plant
of the oviposition plant of the same species was measured.
Potential nectar resources were defined as any forb in bloom at the time of
the survey. The number of blooms per species were counted within the 1 m2
quadrat as well as sampled from eight 1 x ¼ m plots in a 5m radius of the survey
quadrat. The number of shrubs, trees, tree saplings, and tree seedlings were
counted within 10 m of the quadrat. Tree and shrub species present within 20 m of
the quadrat were recorded. The nearest distance from the quadrat to the nearest
forest edge and visible water source as well as the distance to the nearest tree
and nearest shrub were measured. The type of visible water source was
categorized as “none”, “seasonal standing water”, “intermittent stream”, “small
creek”, or “river”. Finally, the slope and aspect were recorded with regard to a 20
m radius of the oviposition and random locations.
2.6 Data Analysis
We delineated available meadow habitat and Mardon use areas by
overlaying our spatial population data on orthophotos in ArcMap GIS. We
established if oviposition and random locations form independent groups among
and within meadows by conducting Discriminant Function Analyses (DFA). We
first conducted a single DFA assessing the differences between both meadow and
location factors (forming 18 groups -nine meadows with two location-types). We
then conducted individual DFAs on a meadow by meadow basis (location-types
forming two groups per meadow analysis). In the meadow specific DFAs, the
11
percent cover and maximum heights of the respective oviposition plant species
were included. Total structure coefficients, here after referred to as “loadings”,
were used to determine which habitat variables contributed to discrimination
between groups. Variables with the highest absolute loading values were
considered the primary descriptors for each canonical axis. As we included over
50 variables per meadow in the DFAs, we considered the first five primary
descriptors as the “most important” variables. We then investigated the next five
primary descriptors to determine if any other variables surfaced more than once
across meadows.
Discriminant Function Analyses were run with SAS statistical software using
the PROC CANDISC procedure. Data were natural-log and arcsine-square root
transformed (for continuous and percent cover data respectively) to achieve
multivariate normality and homogeneous variances. Some of the variables were
not normal, even after appropriate transformations, so we used nonparametric
Mann Whitney U tests to determine significant differences between oviposition and
random locations with regard to the descriptor variables. Bonferroni multiple
comparisons calculated a minimum P-value of P=0.001 (0.05 divided by an
average of 50 variables per site).
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3. Results
Oviposition, habitat, and population surveys were conducted from 31 May
to 1 August, 2007. Survey Meadows range in size from 0.6 ha to 4.3 ha, and
Mardon use areas range from 0.3 ha to 1.3 ha. Except for Grapefern and Flog
Salvage, use areas are smaller than the available open meadow habitat. Minimum
population sizes range from 15 to 313 individuals (Table 1, Appendix C).
We identified nineteen oviposition plant species, including seven sedges
and 12 grasses during the 2007 research season. When including Carex
multicostata, Danthonia californica, Deschampsia cespitosa, and Festuca roemeri
observed only in the 2006 pilot season (at meadows located in Southern Oregon),
there is a total of 23 documented Mardon skipper oviposition plants (Table 2). All
observed oviposition species are native perennials with the exceptions of
nonnative perennial Poa pratensis, and native annual Muhlenbergia filiformis. The
frequency of ovipositions on M. filiformis is low (two observations). Oviposition
species P. pratensis and Festuca idahoensis are most frequently used, both with
56 oviposition observations each (Table 3). However, F. idahoensis was only
present in the two meadows where it was used. Poa pratensis is used for
oviposition across seven meadows, and was present in all meadows except for
Smith Butte (Table 3). Over 75% of ovipositions occur on a single species at Flog
Salvage, Smith Butte, and Midway meadows, indicating graminoid preferences
there. We followed 24 individual female skippers for multiple ovipositions
(Appendix D). Seven of these females clearly switched plant species during
consecutive ovipositions; the remaining seventeen selected a single species.
13
There is no significant separation between oviposition and random locations
when the data are analyzed for discrimination between meadows and locationtype collectively (18 groups including nine meadows, each with two locations),
Figure 3a. However, data group distinctly by meadow factors (Figure 3b),
indicating that the differences among individual meadow habitats overshadow
differences occurring within meadows. Squared canonical correlation (r2) is 88%
and 79% for the first and second discriminating axes respectively. Tree abundance
is the dominant contributing variable to variation on the first axis (variable loading
= -0.883), and percent slope is the dominant contributing variable on the second
axis (variable loading = 0.807).
The meadow by meadow DFAs reveal strong separation between
oviposition and random locations (Figure 4). Squared canonical correlation is over
90% at all meadows except for Smith Butte (r2 = 84%). Primary descriptors are
different at each meadow (Table 4). With regard to the first five primary
descriptors, variables related to graminoid cover and structure are important
across all meadows. Tree variables are important at four meadows. In Flog
Salvage, Muddy, Midway, and Smith Butte specific graminoid species (F.
idahoensis, D. intermedia, and P. pratensis) are important. In these cases the
respective species are the primary oviposition plants (Table 3). The oviposition
plant is the most dominant species in terms of ground cover in the sub-plot (77%
of observations), and in the quadrat (65% of observations). Litter cover and depth
at Grapefern, and soil pH and graminoid richness at Muddy are the first primary
descriptors within those meadows. When we investigate the next five primary
14
descriptors higher graminoid species richness, relative to random locations, are
important at four meadows; greater litter cover and depth appear important at four
meadows; greater vascular plant cover, lower vegetation height and bare ground
cover are each important at three meadows (Figure 5, Appendix E and F).
Oviposition and random locations are significantly different for all primary
discriminating variables (Mann Whitney U tests P<0.001). With few exceptions, all
of the first 5 discriminating variables are significantly different (Table 5). Poa
pratensis cover in the quadrat is not significant at the p=0.05 level in Midway
meadow.
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4. Discussion
Our records of 23 species of oviposition plants, in conjunction with our
observations on individuals alternating host species during consecutive
ovipositions, indicate that Mardon skippers are generalists in terms of oviposition
selection. Other generalist skippers include the Dun (Euphyes vestries), U.S.
federal candidate Dakota (Hesperia dacotae), and Illinois state threatened Ottoe
(Hesperia Ottoe) skippers of which female oviposition and larval feeding on
several graminoid species has been documented (Dana 1997; Shephard 2000).
There are advantages to a generalist life history strategy (Singer et al.
2004; Wee & Singer 2007). Host plant nutrients, microclimate, parasitism and
predation all influence whether or not a particular plant species is used for
oviposition and successfully occupied by larvae (Agosta 2006; Chen et al. 2004;
Fartmann 2006; Lill et al. 2002; Singer & Bernays 2003; Singer et al. 2004;
Smallidge & Leopold 1997). The ability of larvae to utilize multiple species as host
plants reduces restrictions on oviposition selection allowing females to respond to
multi-trophic factors (Doak et al. 2006; Freese et al. 2006; Singer & Bernays
2003). Additionally, larvae may compensate for a poor natal habitat by switching to
more favorable host plants (Albanese et al. 2007; Hellmann 2002).
Pinpointing essential habitat across meadows for the Mardon skipper is
confounded by the vast differences between meadows. The generalist nature of
Mardon likely enables the species to persist in a variety habitat types (Singer et al.
2004; Wee & Singer 2007). Within-meadow analyses reveal that Mardon are
selective with respect to meadow specific habitat components, and allow us to
16
generalize for patterns of habitat use across meadows (Kuussaari et al. 2000;
Singer & Thomas 1996). The variability in primary discriminators between
meadows reflects the response of each Mardon population to their respective
habitats.
Graminoid cover is important at all meadows. Mardon oviposited on larger
graminoids (greater cover) relative to random locations. A higher cover of a host
plant is indicative of a larger larval food resource (Awmack & Leather 2002;
O'Brien et al. 2004). Mardon also select for a greater amount of total vascular plant
cover and a lower amount of bare ground (at three meadows). Bare ground
restricts movement of larvae and exposes them to predation, where a greater
amount of vegetative cover may provide protection (Doak 2000).
The percent cover of specific graminoid species is important at seven of the
nine meadows; F. idahoensis, F. rubra, P. pratensis, A. thurberiana, and D.
intermedia. Additionally, 77% of oviposition plant species had the most amount of
cover relative other species present in the egg vicinity (sub-plot), and 65% in the
local vicinity (quadrat). These results indicate that Mardon may prefer some
graminoid species over others. However, every plant species has an inherent
structure and distinct invertebrate community (Reid & Hochuli 2007). Strong within
meadow preferences for specific species may reflect a selection for host plant
architecture, chemistry, or nutrient value (Reid & Hochuli 2007; Talsma et al. 2008;
Wee & Singer 2007) rather than a species specific dependency.
Oviposition plant leaf density is important at 7A, Cave Creek, Flog Salvage
and Smith Butte meadows. Additionally, graminoid height is important at four
17
meadows and maximum vegetation height is important at three meadows. This
result enforces that vegetation height and graminoid structure (height, leaf size, &
density) is important in the local area (quadrat), and egg vicinity (sub-plot), and is
an important characteristic of the oviposition plant. The structure of the host plant
and the surrounding vegetation directly influences the microclimate (humidity,
temperature, solar exposure) important for egg and larval development; and
affords protection from predation and parasitism (Awmack & Leather 2002; Freese
et al. 2006). The silver spotted skipper (Hesperia comma), and duke of burgundy
butterfly (Hamearis lucina), have specific structural requirements for their larval
host plants (Agosta 2008; Davies et al. 2005; New 1997; Thomas & Jones 1993).
At four sites graminoid species richness and litter factors appear in the first
ten primary discriminating variables. Species richness is higher in oviposition
locations relative to random locations at all sites except Peterson Prairie. Insect
herbivore species richness increases with plant species richness (Haddad et al.
2001; Panzer & Schwartz 1998; Reid & Hochuli 2007) as there is a greater
availability of alternate vegetative resources and structure (Haddad et al. 2001).
Plant species richness is also indicative of greater soil nutrients, which in turn
affect the diets of insect herbivores (Haddad et al. 2001; Ockinger et al. 2006).
Greater litter cover and depth likely offer protection during egg and larval
life stages. The fungal pathogen (Entomophaga maimaiga), specific to
Lepidopteran species, rarely occurs in species that inhabit litter layers during the
larval state (Hajek et al. 2000). Additionally, forest dwelling Lepidoptera often
pupate under litter layers on the ground (Dugdale 1996), where they are less
18
exposed to extreme weather conditions. During our 2006 pilot season we found
Mardon larvae to be active just prior to the first snow fall, which is an indication
that they overwinter in this state (Beyer & Black 2006). Therefore, litter may have
an insolation benefit to skippers, protecting them from extreme temperatures
during early life stages.
The data from five of nine meadows indicate that tree factors are important
to oviposition selection. At these meadows tree canopy cover and tree abundance
are negatively associated with oviposition locations indicating a preference for low
cover and low tree abundance. Distance to forest edge and distance to nearest
tree are positively associated with oviposition locations, indicating a preference for
larger distances from trees. A high amount of tree and shrub cover reduces solar
insolation and shading of habitat creates a cooler environment. As butterflies are
ectotherms, they are physiologically limited to daylight hours when temperatures
are high enough to enable butterfly mobility (Davies et al. 2005; Doak et al. 2006;
Freese et al. 2006). Additionally, egg-laying rates are temperature dependent
(Davies et al. 2006). The fact that Mardon skippers lay eggs singly places further
time restrictions for selecting suitable host plants, as there is a greater time
investment per egg (Courtney 1984).
Graminoid communities change in response to forest proximity due to
shading effects. Encroaching conifers alter the composition of soil creating
favorable conditions for tree seedling establishment while making soils unsuitable
for meadow-specific vegetation (Griffiths et al. 2005). Forest encroachment also
reduces the soil microbial communities important to nitrogen fixation of meadow
19
grasses and forbs (Griffiths et al. 2005), and likely reduces the nutrient content of
larval food plants. It is likely that Mardon skippers select for more exposed
oviposition locations because they are selecting for exposed open meadow
habitat, and the graminoid communities that are associated with it.
4.1 Conservation implications
Our results suggest that preserving Mardon skippers in montane meadow
habitats will require active management to control forest encroachment, as well as
maintain meadow specific graminoid communities and structure important to their
respective Mardon populations. The removal and alteration of the natural
disturbance regimes (such as fire suppression) that once maintained low conifer
seedling establishment rates, has led to the loss and degradation of forestmeadow ecosystems (Coop & Givnish 2007; Norman & Taylor 2005). Forest
encroachment not only reduces the amount of open habitat but closes off corridors
between meadows reducing butterfly dispersal (Roland & Matter 2007).
All of the Mardon populations in this study are isolated by thick forest
barriers, and exist in low numbers. A better understanding of the feasibility and the
effects of reconnecting neighboring Mardon populations that have been isolated by
forest encroachment is warranted (Bergman 1999; Dennis et al. 2006). Careful
consideration should be given to the genetic and behavioral implications of local
habitat preferences (Kuussaari et al. 2000; Moreau et al. 2008) as Mardon have
displayed meadow specific selectivity in our research.
There is a fine balance between incorporating enough disturbance to
maintain meadow ecosystems and too much disturbance causing habitat
20
degradation and butterfly mortality (Schultz et al. 2008). Livestock grazing
adversely impacts butterfly populations by altering plant community composition
(Stoner & Joern 2004) and trampling during immobile life stages (egg, larvae,
pupae) or during cool temperatures when adult movement is restricted (Warren
1993a, b). Over-grazing can be detrimental by stripping habitat of vegetation,
removing adult nectar resources, and introducing invasive weeds (Hayes & Holl
2003). However, light-rotational grazing, mowing, and burning can maintain
vegetation heights and habitat heterogeneity favorable to butterflies (Ravenscroft
1994; Vogel et al. 2007). The silver spotted skipper has not only shown a positive
response to moderate grazing, but depends on it to maintain the structure of its
host plant (Davies et al. 2005; Thomas & Jones 1993). Similarly, to achieve the
benefits of restorative burning, fire treatments must be carefully prescribed to
prevent local extinction (Schultz & Crone 1998; Vogel et al. 2007).
Understanding the effects of non-native invasive species on Mardon
skippers is critical. All oviposition plants in this study are native species, except for
P. pratensis, and all but one species are perennials. When present in meadows F.
idahoensis is clearly preferred over other species, which supports former
conceptions about the importance of Festuca grasses to this species (Black &
Vaughan 2005; Potter et al. 2002). Poa pratensis is the most widely used
oviposition plant across meadows, but is generally not heavily selected for within
meadows (Table 4). Poa pratensis has been correlated with increased
abundances and invertebrate species richness in other studies (Reid & Hochuli
2007), and may be selected for by skippers because it is structurally similar to
21
other native oviposition plants at our research meadows. Non-native grasses that
do not fit the structural and nutritive requirements of Mardon may, however, be
detrimental to populations. Invasive plants tend to out-compete native
communities, and negatively impact grassland ecosystems by homogenizing the
habitat (Kolb et al. 2002; Possley & Maschinski 2006).
Continued research into larval preference and larval survivorship is
essential to understand host plant characteristics vital to larval Mardon skippers.
Our results indicate that graminoid abundance and architecture are key
discriminators for oviposition selection. A more extensive and refined look into the
characteristics of oviposition plants and surrounding vegetation is warranted.
Understanding adult habitat use, movement within and dispersal from habitat
patches is also essential to developing effective conservation strategies (Kuefler &
Haddad 2006; Ries & Debinski 2001; Schtickzelle et al. 2007; Schultz & Crone
2008; Turchin et al. 1991).
Anthropogenic land use has shaped meadow and grassland habitats
globally (Crawford & Hall 1997; Davies et al. 2005; Louy et al. 2007; Ockinger et
al. 2006). Relative to all other butterfly families, knowledge of the basic life history
requirements of grass skippers is poor (Wahlberg et al. 2005). To curb skipper
declines, knowledge about their biology must be expanded. Our research has
taken the first step in understanding what influences oviposition behavior of the
Mardon skipper. It is clear from our work that Mardon skippers select for open
habitats, and specific vegetative structure. Though Mardon are not regionally
selective for specific grass species, they do exhibit oviposition plant specificity
22
within localities. The vast differences between Mardon meadows, and the variation
in selectivity among meadows, indicate that there are no simple solutions to
conserving this butterfly. A more complete understanding of Mardon biology and
resource use across the habitats of extant populations is necessary before broad
and regional generalizations can be reliably made.
23
REFERENCES
Agosta, S. J. 2006. On ecological fitting, plant-insect associations, herbivore host
shifts, and host plant selection. Oikos 114:556-564.
Agosta, S. J. 2008. Fitness consequences of host use in the field: temporal
variation in performance and a life history tradeoff in the moth Rothschildia
lebeau (Saturniidae). Oecologia 157:69-82.
Albanese, G., P. Vickery, and P. Sievert. 2007a. Microhabitat use by larvae and
females of a rare barrens butterfly, frosted elfin (Callophrys irus ). Journal of
Insect Conservation.
Albanese, G., P. D. Vickery, and P. R. Sievert. 2007b. Habitat characteristics of
adult frosted elfins (Callophrys irus) in sandplain communities of
southeastern Massachusetts, USA. Biological Conservation 136:53-64.
Anthes, N., T. Fartmann, and G. Hermann. 2008. The Duke of Burgundy butterfly
and its dukedom: larval niche variation in Hamearis lucina across Central
Europe. Journal of Insect Conservation 12:3-14.
Anthes, N., T. Fartmann, G. Hermann, and G. Kaule. 2003. Combining larval
habitat quality and metapopulation structure-the key for successful
management of pre-alpine Euphydryas aurina colonies. Journal of Insect
Conservation 7:175-185.
Awmack, C. S., and S. R. Leather. 2002. Host plant quality and fecundity in
herbivorous insects. Annual Review of Entomology 47:817-844.
Bergman, K. O. 1999. Habitat utilization by Lopinga achine (Nymphalidae :
Satyrinae) larvae and ovipositing females: implications for conservation.
Biological Conservation 88:69-74.
Bergman, K. O., J. Askling, O. Ekberg, H. Ignell, H. Wahlman, and P. Milberg.
2004. Landscape effects on butterfly assemblages in an agricultural region.
Ecography 27:619-628.
Brown, K. S., and A. V. L. Freitas. 2000. Atlantic forest butterflies: Indicators for
landscape conservation. Biotropica 32:934-956.
Calabrese, J. M., L. Ries, S. F. Matter, D. M. Debinski, J. N. Auckland, J. Roland,
and W. F. Fagan. 2008. Reproductive asynchrony in natural butterfly
populations and its consequences for female matelessness. Journal of
Animal Ecology 77:746-756.
Chen, L. L., and Z. S. Lin. 2008. The effect of habitat destruction on
metapopulations with the Allee-like effect: A study case of Yancheng in
Jiangsu Province, China. Ecological Modelling 213:356-364.
Chen, Y. Z., L. Lin, C. W. Wang, C. C. Yeh, and S. Y. Hwang. 2004. Response of
two Pieris (Lepidoptera : Pieridae) species to fertilization of a host plant.
Zoological Studies 43:778-786.
Courtney, S. P. 1984. The evolution of egg elustering by butterflies and other
insects. The American Naturalist 123:276-281.
Davies, Z. G., R. J. Wilson, T. M. Brereton, and C. D. Thomas. 2005. The reexpansion and improving status of the silver-spotted skipper butterfly
(Hesperia comma) in Britain: a metapopulation success story. Biological
24
Davies, Z. G., R. J. Wilson, S. Coles, and C. D. Thomas. 2006. Changing habitat
associations of a thermally constrained species, the silver-spotted skipper
butterfly, in response to climate warming. Journal of Animal Ecology
75:247-256.
Davis, J. D., D. M. Debinski, and B. J. Danielson. 2007. Local and landscape
effects on the butterfly community in fragmented Midwest USA prairie
habitats. Landscape Ecology 22:1341-1354.
Dennis, R. L. H., T. G. Shreeve, and H. Van Dyck. 2006. Habitats and resources:
The need for a resource-based definition to conserve butterflies.
Biodiversity and Conservation 15:1943-1966.
Doak, P. 2000. Population consequences of restricted dispersal for an insect
herbivore in a subdivided habitat. Ecology 81:1828-1841.
Doak, P., P. Kareiva, and J. Kingsolver. 2006. Fitness consequences of choosy
oviposition for a time-limited butterfly. Ecology 87:395-408.
Dugdale, J. S. 1996. Natural history and identification of litter-feeding Lepidoptera
larvae (Insecta) in beech forests, Orongorongo Valley, New Zealand, with
especial reference to the diet of mice (Mus musculus). Journal of the Royal
Society of New Zealand 26:251-274.
Eastwood, R., and J. M. Hughes. 2003. Molecular phylogeny and evolutionary
biology of Acrodipsas (Lepidoptera : Lycaenidae). Molecular Phylogenetics
and Evolution 27:93-102.
Ellis, S. 2003. Habitat quality and management for the northern brown argus
butterfly Aricia artaxerxes (Lepidoptera : Lycaenidae) in North East
England. Biological Conservation 113:285-294.
Fartmann, T. 2006. Oviposition preferences, adjacency of old woodland and
isolation explain the distribution of the Duke of Burgundy butterfly
(Hamearis lucina) in calcareous grasslands in central Germany. Annales
Zoologici Fennici 43:335-347.
Fownes, S., and J. Roland. 2002. Effects of meadow suitability on female
behaviour in the alpine butterfly Parnassius smintheus. Ecological
Entomology 27:457-466.
Fox, R., T. B. Asher, D. Roy, and M. Warren. 2006. The state of butterflies in
Britain and Ireland. J Insect Conserv.
Fred, M. S., R. B. O'Hara, and J. E. Brommer. 2006. Consequences of the spatial
configuration of resources for the distribution and dynamics of the
endangered Parnassius apollo butterfly. Biological Conservation 130:183192.
Freese, A., J. Benes, R. Boz, O. Cizek, M. Dolek, A. Geyer, P. Gros, M. Konvicka,
A. Liegl, and C. Stettmer. 2006. Habitat use of the endangered butterfly
Euphydryas maturna and forestry in Central Europe. Animal Conservation
9:388-397.
Griffiths, R., M. Madritch, and A. Swanson. 2005. Conifer invasion of forest
meadows transforms soil characteristics in the Pacific Northwest. Forest
Ecology and Management 208:347-358.
25
Grundel, R., and N. B. Pavlovic. 2007. Resource availability, matrix quality,
microclimate, and spatial pattern as predictors of patch use by the Karner
blue butterfly. Biological Conservation 135:135-144.
Grundel, R., N. B. Pavlovic, and C. L. Sulzman. 1998. Habitat use by the
endangered Karner blue butterfly in oak woodlands: the influence of canopy
cover. Biological Conservation 85:47-53.
Haddad, N. M., D. Tilman, J. Haarstad, M. Ritchie, and J. M. H. Knops. 2001.
Contrasting effects of plant richness and composition on insect
communities: A field experiment. American Naturalist 158:17-35.
Hajek, A. E., L. Butler, J. K. Liebherr, and M. H. Wheeler. 2000. Risk of infection
by the fungal pathogen Entomophaga maimaiga among Lepidoptera on the
forest floor. Environmental Entomology 29:645-650.
Hayes, G. F., and K. D. Holl. 2003. Cattle grazing impacts on annual forbs and
vegetation composition of mesic grasslands in California. Conservation
Biology 17:1694-1702.
Hellmann, J. 2002. The effect of an environmental change on mobile butterfly
larvae and nutritional quality of their hosts. Journal of Animal Ecology
71:925-936.
Kolb, A., P. Alpert, D. Enters, and C. Holzapfel. 2002. Patterns of invasion within a
grassland community. Journal of Ecology 90:871-881.
Kuefler, D., and N. M. Haddad. 2006. Local versus landscape determinants of
butterfly movement behaviors. Ecography 29:549-560.
Kuhrt, U., J. Samietz, and S. Dorn. 2006. Thermal response in adult codling moth.
Physiological Entomology 31:80-88.
Kuussaari, M., I. Saccheri, M. Camara, and I. Hanski. 1998. Allee effect and
population dynamics in the Glanville fritillary butterfly. OIKOS 82:384-392.
Kuussaari, M., M. Singer, and I. Hanski. 2000. Local specialization and landscapelevel influence on host use in an herbivorous insect. Ecology 81:2177-2187.
Leung, Y., and J. L. Marion. 2000. Recreation impacts and management in
wilderness: a state of-knowledge review. . USDA Forest Service
Proceedings 25.
Lill, J. T., R. J. Marquis, and R. E. Ricklefs. 2002. Host plants influence parasitism
of forest caterpillars. Nature 417:170-173.
Liu, W. H., Y. F. Wang, and R. M. Xu. 2006. Habitat utilization by ovipositing
females and larvae of the Marsh fritillary (Euphydryas aurinia) in a mosaic
of meadows and croplands. Journal of Insect Conservation 10:351-360.
Mizumoto, M., and F. Nakasuji. 2007. Egg size manipulation in the migrant
skipper, Parnara guttata guttata (Lepidoptera : Hesperiidae), in response to
different host plants. Population Ecology 49:135-140.
Moreau, J., J. Rahme, B. Benrey, and D. Thiery. 2008. Larval host plant origin
modifies the adult oviposition preference of the female European grapevine
moth Lobesia botrana. Naturwissenschaften 95:317-324.
Myers, J. H. 1985. Effect of physiological condition of the host plant on the
ovipositional choice of the Cabbage white butterfly, Pieris rapae. Journal of
Animal Ecology 54:193-204.
26
New, T. R., editor. 1997. Butterfly Conservation. Oxford University Press Australia,
Melbourne.
Norman, S. P., and A. H. Taylor. 2005. Pine forest expansion along a forestmeadow ecotone in northeastern California, USA. Forest Ecology and
Management 215:51-68.
Noss, R. F., E. T. L. III, and J. M. Scott 1995. Endangered ecosystems of the
United States: A preliminary assessment of loss and degradation. National
Biological Service, Washington, D. C.
O'Brien, D. M., C. L. Boggs, and M. L. Fogel. 2004. Making eggs from nectar: the
role of life history and dietary carbon turnover in butterfly reproductive
resource allocation. Oikos 105:279-291.
Ockinger, E., O. Hammarstedt, S. G. Nilsson, and H. K. Smith. 2006. The
relationship between local extinctions of grassland butterflies and increased
soil nitrogen levels. Biological Conservation 128:564-573.
Oostermeijer, J. G. B., and C. A. M. van Swaay. 1998. The relationship between
butterflies and environmental indicator values: a tool for conservation in a
changing landscape. Biological Conservation 86:271-280.
Panzer, R., and M. W. Schwartz. 1998. Effectiveness of a vegetation-based
approach to insect conservation. Conservation Biology 12:693-702.
Portugal, A. H. A., and J. R. Trigo. 2005. Similarity of cuticular lipids between a
caterpillar and its host plant: A way to make prey undetectable for predatory
ants? Journal of Chemical Ecology 31:2551-2561.
Possley, J., and J. Maschinski. 2006. Competitive effects of the invasive grass
Rhynchelytrum repens (Willd.) CE Hubb. on pine rockland vegetation.
Natural Areas Journal 26:391-395.
Ravenscroft, N. O. M. 1994. The Ecology of the Chequered skipper butterfly
Carterocephalus palaemon in Scotland .1. Microhabitat. Journal of Applied
Ecology 31:613-622.
Reid, A. M., and D. F. Hochuli. 2007. Grassland invertebrate assemblages in
managed landscapes: Effect of host plant and microhabitat architecture.
Austral Ecology 32:708-718.
Ries, L., and D. M. Debinski. 2001. Butterfly responses to habitat edges in the
highly fragmented prairies of Central Iowa. Journal of Animal Ecology
70:840-852.
Roland, J., and S. F. Matter. 2007. Encroaching forests decouple alpine butterfly
population dynamics. Proceedings of the National Academy of Sciences
104:13702-13704.
Schtickzelle, N., C. Turlure, and M. Baguette. 2007. Grazing management impacts
on the viability of the threatened bog fritillary butterfly Proclossiana
eunomia. Biological Conservation 136:651-660.
Schultz, C. B., and E. E. Crone. 1998. Burning prairie to restore butterfly habitat: A
modeling approach to management tradeoffs for the Fender's blue.
Restoration Ecology 6:244-252.
Schultz, C. B., and E. E. Crone. 2008. Note: Using ecological theory to advance
butterfly conservation. Israel Journal of Ecology & Evolution 54:63-68.
27
Schweiger, O., C. F. Dormann, D. Bailey, and M. Frenzel. 2006. Occurrence
pattern of Pararge aegeria (Lepidoptera : Nymphalidae) with respect to
local habitat suitability, climate and landscape structure. Landscape
Ecology 21:989-1001.
Singer, M. C., and C. D. Thomas. 1996. Evolutionary responses of a butterfly
metapopulation to human- and climate-caused environmental variation.
American Naturalist 148:S9-S39.
Singer, M. S., and E. A. Bernays. 2003. Understanding omnivory needs a
behavioral perspective. Ecology 84:2532-2537.
Singer, M. S., D. Rodrigues, J. O. Stireman, and Y. Carriere. 2004. Roles of food
quality and enemy-free space in host use by a generalist insect herbivore.
Ecology 85:2747-2753.
Smallidge, P. J., and D. J. Leopold. 1997. Vegetation management for the
maintenance and conservation of butterfly habitats in temperate humandominated landscapes. Landscape and Urban Planning 38:259-280.
Stoner, K. J. L., and A. Joern. 2004. Landscape vs. local habitat scale influences
to insect communities from tallgrass prairie remnants. Ecological
Applications 14:1306-1320.
Stuart, B. W. 1999. Cars, cows, and checkerspot butterflies: Nitrogen deposition
and management of nutrient-poor grasslands for a threatened species.
Conservation Biology 13:1476-1486.
Talsma, J. H. R., A. Biere, J. A. Harvey, and S. van Nouhuys. 2008. Oviposition
cues for a specialist butterfly-plant chemistry and size. Journal of Chemical
Ecology 34:1202-1212.
Thomas, C. D., E. J. Bodsworth, R. J. Wilson, A. D. simmons, Z. G. Davies, M.
Musche, and L. Conradt. 2001. Ecological and evolutionary processes at
expanding range margins. Nature 411:577-581.
Thomas, C. D., and T. M. Jones. 1993. Partial recovery of a skipper butterfly
(Hesperia comma) from population refuges- lessons for conservation in a
fragmented landscape. Journal of Animal Ecology 62:472-481.
Thomas, J. A., M. G. Morris, and C. Hambler. 1994. Patterns, mechanisms, and
rates of extinction among invertebrates in the United-Kingdom.
Philosophical Transactions of the Royal Society of London Series BBiological Sciences 344:47-54.
Trombulak, S. C., and C. A. Frissell. 2000. Review of ecological effects of roads
on terrestrial and aquatic communities. Conservation Biology 14:18-30.
Turchin, P., F. J. Odendaal, and M. D. Rausher. 1991. Quantifying insect
movement in the field. Environmental Entomology 20:955-963.
Vogel, J. A., D. M. Debinski, R. R. Koford, and J. R. Miller. 2007. Butterfly
responses to prairie restoration through fire and grazing. Biological
Wallisdevries, M. F., and C. A. M. Van Swaay. 2006. Global warming and excess
nitrogen may induce butterfly decline by microclimatic cooling. Global
Change Biology 12:1620-1626.
28
Warren, M. S. 1993a. A review of butterfly conservation in Central Southern Britain
.1. Protection, evaluation and extinction on prime sites. Biological
Warren, M. S. 1993b. A review of butterfly conservation in Central Southern Britain
.2. Site management and habitat selection of key species. Biological
Wee, B., and M. C. Singer. 2007. Variation among individual butterflies along a
generalist-specialist axis: no support for the 'neural constraint' hypothesis.
Ecological Entomology 32:257-261.
29
Table 1: Meadow Descriptions
Description of 9 Mardon skipper research meadows in the Gifford Pinchot National Forest, Washington State. All
meadows may be exposed to elk grazing.
Meadow
Cave Creek
Elevation (m) Aspect
850
Flat
Habitat Size (ha)
Available
Min Use
Area
Area
2
1
Total Days Surveys were Conducted
Minimum Flight Period
Oviposition
6
Habitat
10
Population
4
Population
Size
56
From
May 31
To
June 23
Impacts/Management
Past & current partial
grazing, fenced, road,
noxious weeds removal
management
Past grazing, fenced,
road
915
Flat
4.3
1.3
3
5
3
34
June 8
June 28
Lost
975
Flat
1.5
1.1
1
5
3
15
June 18
June 29
Grazing
Flog Salvage
1190
Flat
<0.8
<0.8
3
4
3
23
June 21
June 30
Logged and seeded
historically, road
Midway
1280
North
2
1.3
2
3
3
54
July 8
Aug 1
Recreation, horse
watering & grazing, road,
campground
Smith Butte
1295
South
0.5
0.3
1
3
2
38
July 1
July 3
Light grazing
Muddy
1340
Mixed
1.5
0.9
2
5
3
91
July 12
Aug 2
Hiking trail, Horse back
riding & grazing
7A
1430
West
0.9
0.6
2
3
3
50
July 8
Aug 1
Grapefern
1430
West
0.6
0.6
1
3
3
313
July 9
Aug 1
riding & grazing, Conifer
encroachment
management
riding & grazing, Conifer
sapling removal
30
Peterson
Table 2: Oviposition Plant Species
Observed oviposition plant species in the 2007 research and 2006 pilot seasons. Species are native perennials unless
otherwise noted. A=annual, NI=non-native invasive, 2006 = only observed in the pilot study.
Species
Common Name
Alternate names
Long-stolon Sedge
Short Stem Sedge
Woodrush Sedge
Hall's Sedge
Hood's Sedge
Fragile Sheathed Sedge
Manyrib Sedge
Meadow Sedge
C. pensylvanica
C. brevicaulis, C. rossii
Thurber Bent
California Brome
Bluejoint
Tufted Hairgrass
Timber Oatgrass
California Oatgrass
One-spike Oatgrass
Idaho Fescue
Red Fescue
Roemer's Fescue
Meadow Barley
Pullup Muhly
Kentucky Bluegrass
Squirreltail
Western Needlegrass
A. humilis, Podagrostis thurberiana
B. marginatus
Sedges
31
Carex inops
Carex deflexa
Carex luzulina
Carex halliana
Carex hoodii
Carex fracta
Carex multicostata 2006
Carex praticola
Grasses
Agrostis thurberiana
Bromus carinatus
Calamagrostis canadensis
Deschampsia cespitosa 2006
Danthonia intermedia
Danthonia californica 2006
Danthonia unispicata
Festuca idahoensis
Festuca rubra
Festuca roemeri 2006
Hordeum brachyantherum
Muhlenbergia filiformis A
Poa pratensis NI
Elymus elymoides
Stipa occidentalis
Danthonia canadensis
D. americana
M. idahoensis, M. simplex
Sitanion hystrix
Achnatherum occidentalis
Table 3: Oviposition Frequency By Plant Species
The frequency of ovipositions per graminoid species at each of the 9 research meadows in 2007.
32
Oviposition Plant Species
7A
Cave Creek
Festuca idahoensis
Poa pratensis
3
8
Danthonia intermedia
10
Carex inops
2
Festuca rubra
1
17
Carex deflexa
1
Carex fracta
7
Carex praticola
Carex hoodii
3
Danthonia unispicata
3
Stipa occidentalis
1
Carex halliana
Carex species
Bromus carinatus
Muhlenbergia filiformis
1
Unknown Grass
1
Calamagrostis canadensis
Carex luzulina
1
Elymus elymoides
Hordeum brachyantherum
Unknown Sedge
- Species present at site but not used for oviposition
Flog
Salvage
27
3
Lost
Midway
Muddy
2
1
14
5
23
1
4
15
-
11
-
11
3
1
3
1
-
-
Peterson
Prairie
Grapefern
2
2
3
1
13
1
2
8
2
1
1
5
4
-
-
1
1
1
1
1
1
-
-
1
2
1
Smith Butte Total
29
56
56
27
19
1
19
16
12
11
10
9
8
8
4
3
2
2
2
1
1
1
1
1
Table 4: Within Meadow Primary Descriptors
Top 5 primary discriminating variables in meadow by meadow Discriminant Function
Analysis. Total structure coefficients, means ± standard deviations for oviposition and
random locations, and unit of measure given. Corresponds with Figure 4.
Location
Oviposition
Random
8.5 ± 7.3
18 ± 10.1
34.8 ± 14.0 20.5 ± 16.2
89.1 ± 19.6 64.9 ± 30.0
2.2 ± 7.0
1.5 ± 0.6
5.5 ± 4.6
13.3 ± 11.4
Discriminatory Axis Variable
Tree Abundance
Graminoid Cover (Egg Vicinity)
Distance To Visible Water Source1
1
Oviposition Plant Leaf Density
Tree Canopy Cover 1
Variable
Loadings
-0.486
0.460
0.440
0.439
-0.433
Maximum Plant Height
Distance To Nearest Tree
Festuca rubra Cover (Egg Vicinity)
Tree Canopy Cover
0.657
-0.588
0.539
-0.474
0.455
Flog Salvage
Festuca idahoensis Cover (Egg Vicinity)
Oviposition Plant Footprint
Graminoid Cover (Quadrat)
0.783
0.772
0.709
0.679
0.614
Grapefern
Litter Depth
Tree Abundance
0.670
0.608
0.568
0.541
-0.486
2.1 ± 0.9
48.9 ± 12.8
82.5 ± 10.9
50.8 ± 17.1
3.0 ± 3.9
0.8 ± 0.8
27 ± 15.5
53.8 ± 28.0
27.5 ± 19.2
16.3 ± 20.9
cm
%
%
%
count
Vascular Plant Cover (Quadrat)
Poa pratensis Cover (Quadrat)
1
Bare Ground Cover
-0.600
-0.540
-0.479
-0.476
0.471
54.7 ± 18.2
53.2 ± 18.3
68.5 ± 16.7
16.0 ± 9.9
10.3 ± 6.5
29.1 ± 19.2
32.3 ± 16.3
51.3 ± 16.9
8.4 ± 5.4
28.3 ± 22.9
%
%
%
%
%
Muddy
Soil Ph
Graminoid Species Richness (Quadrat)1
Danthonia intermedia Height (Quadrat)1
Danthonia intermedia Cover (Quadrat)
Agrostis thurberiana Cover (Quadrat)1
-0.477
0.474
0.467
0.441
0.429
6.1 ± 0.3
6.4 ± 1.4
34.5 ± 7.0
23.7 ± 11.9
22.4 ± 26.1
6.4 ± 0.3
4.9 ± 1.5
33.9 ± 8.5
23.0 ± 21.1
11.7 ± 5.6
scale
count
cm
%
%
Midway
Poa pratensis Height (Quadrat)
Poa pratensis Cover (Quadrat) 2
Graminoid Cover (Quadrat) 1
1
Bare Ground Cover
Flower Abundance (5m radius) 1
-0.456
0.364
0.354
-0.351
0.318
31.3 ± 9.1
28.8 ± 24.1
49 ± 20.5
5.4 ± 6.7
12.3 ± 8.9
41.1 ± 11.2
15.1 ± 10.8
36.2 ± 14.6
15.8 ± 20.0
7.7 ± 6.1
cm
%
%
%
count
0.520
0.509
-0.501
0.494
0.461
54.2 ± 11.8
39.1 ± 12.5
0.4 ± 1.1
17 ± 8.8
2.1 ± 0.03
37.7 ± 17.0
23.5 ± 15.8
7.3 ± 10.9
9.8 ± 5.3
2.0 ± 0.04
meters
meters
%
%
scale
Meadow
7A
Cave Creek
3
Litter Cover
Lost
3
Peterson Prairie Distance To Forest Edge
Distance To Nearest Tree
Tree Canopy Cover
Poa pratensis Cover (Quadrat)
Graminoid Cover (Quadrat)1
Smith Butte3
52.8 ± 10.8
1.8 ± 0.7
4.7 ± 4.5
19.1 ± 18.4
16.4 ± 17.2
cm
category (1-4)
meters
%
%
34.3 ± 20.6 12.2 ± 12.2
212.3 ± 168.1 30.7 ± 29.1
2.8 ± 0.5
1.6 ± 0.7
42.6 ± 20.1 16.5 ± 10.1
34.7 ± 11.9
20.3 ± 8.0
%
cm2
category (1-4)
%
%
38.4 ± 6.2
2.7 ± 0.7
1.5 ± 1.6
32.2 ± 12.6
5.4 ± 7.4
-0.694
2.2 ± 0.4
1.5 ± 0.5
Festuca idahoensis Cover (Egg Vicinity)
24.0 ± 13.9 12.2 ± 9.6
-0.686
Festuca idahoensis Height (Egg Vicinity)
25.2 ± 31.4
14.8 ± 3.3
-0.649
-0.505
24.3 ± 13.7
16.4 ± 9.8
Festuca idahoensis Cover (Quadrat) 1
Oviposition Plant Footprint
-0.503
76.4 ± 61.3 40.9 ± 116.0
1 Mann-Whitney U test not significant after correcting for multiple comparisons
2 Mann-Whitney U test not significant at the 0.05 level
3 Class means indicate that oviposition locations are negatively correlated with the canonical function
33
Unit of Measure
count
%
meters
category (1-4)
%
category (1-4)
%
cm
%
cm2
Figure 1: Research Meadows
Nine research meadows located in the Gifford Pinchot National Forest in
Washington State, USA
34
Figure 2: Quadrat and Sub-plot Diagram
1 m2 quadrat and 0.1 m2 sub-plot (egg vicinity)
1m
1m
1/10 m
egg
35
Figure 3: Among Meadow Discriminate Analysis
DFA results on n=540 observations by meadow and location collectively, 18
groups (9 meadows two location-types each). Data symbolized by Location-type
(a) and by meadow (b). Squared canonical correlation = r2.
6
(a)
Percent Slope
2
Total Structure Coefficient = 0.807, r = 0.79, eigenvalue = 3.9
4
2
0
-2
-4
Oviposition
Random
-6
-12
-10
-8
-6
-4
-2
0
2
4
6
-6
-4
-2
0
2
4
6
6
(b)
4
2
0
7A
Cave Creek
Flog Salvage
Grapefern
Lost
Midway
Muddy
Peterson Prairie
Smith Butte
-2
-4
-6
-12
-10
-8
Tree Abundance
2
Total Structure Coefficient = -0.88, r = 0.88, eigenvalue = 7.4
36
Figure 4: Within Meadow Discriminant Function Analyses
Meadow by meadow DFA grouped by location. Oviposition locations represented in
black random locations in white. Percent of discriminant variation explained by
variables = r2. Variables corresponding to X-axes given in Table 4.
Grapefern
7A
2
r : 0.98
14
12
Midway Meadow
2
r : 0.99
10
r2: 0.93
8
8
6
10
6
8
4
6
4
4
0
0
-12 -10 -8 -6 -4 -2 0
No. Observations
2
2
2
2
4
6
8 10 12
0
-10
-5
5
10
-6
8
2
2
r : 0.99
-4
0
2
4
r2: 0.91
r : 0.95
8
-2
Peterson Prairie
Lost Meadow
Cave Creek
10
0
6
8
6
6
4
4
4
2
2
2
0
0
-6
-4
-2
0
2
4
6
8
-4
-2
0
2
4
6
-6
-4
-2
Muddy Meadow
Flog Salvage
2
r : 0.92
7
0
-6
2
4
Smith Butte
10
2
r2: 0.84
r : 0.99
8
0
8
No. Observations
6
5
4
3
2
6
6
4
4
2
2
1
0
0
-6
-4
-2
0
2
4
6
0
-20
20
Canonical Coefficient
37
-4
-2
0
2
4
Figure 5: Across Meadow Trends
Across meadow trends showing variables important to discriminating axes in Figure 4. First five primary descriptors in
black, next five primary descriptors in gray. Q = 1 m2 quadrat and E = egg vicinity (0.1 m2 subplot).
10
No. Meadows
8
6
4
38
Graminoid
Oviposition
Plant
Tree/Shading
Litter
Other
Vegetation Height (Q)
Vasscular cover (Q)
Bare ground cover (Q)
Depth (E)
Litter cover (Q)
Forest edge
Canopy Cover
Tree abundance
Footprint
Leaf density
Richness (E)
Richness (Q)
Species height (E)
Species height (Q)
Total cover (E)
Total cover (Q)
Species cover (E)
Species cover (Q)
0
Nearest tree
2
APPENDICES
A. Examples of Global At-Risk Skippers ...........................................................40
B. Measured Habitat Variables ...........................................................................44
C. Population Counts ..........................................................................................45
D. Multiple Consecutive Ovipositions ...............................................................46
E. Within Meadow DFA Variable Descriptions..................................................47
F. Within Meadow DFA λ, r2, and Class Mean Values ......................................56
G. Temperature and Relative Humidity..............................................................57
Appendix A: Examples of Global At-Risk Skippers
40
In the grass skipper subfamily (Hesperiinae) Y= yes, N=no. Generalist larval diets include more than one plant species
Species
Common Name Status
Larval Diet Hesperiinae?
Europe
Chequered
Priority (Scotland), Endangered
Specialist
N
Carterocephalus palaemon
(Japan)
palaemon
Dingy
Priority species (UK, Ireland)
Generalist
N
Erynnis tages
Silver spotted
Rare, Protected (UK)
Specialist
Y
Hesperia comma comma
Cinquefoil
Threatened, Vulnerable, Declining
Specialist
N
Pyrgus cirsii
(Throughout Europe)
Grizzled
Priority species (UK, Ireland)
Generalist
N
Pyrgus malvae
Lulworth
Protected (UK)
Specialist
Y
Thymelicus acteon
Austrailia
Cynone grass
Vulnerable
Generalist
N
Anisynta cynone cynone
White veined
Vulnerable
Generalist
N
Herimosa albovenata albovenata
grass
Vulnerable
Generalist
N
Hesperilla chrysotricha cyclospila Chrysotricha
sedge
Yellowish sedge Endangered
Generalist
N
Hesperilla donnysa donnysa
Flame sedge
Vulnerable
Generalist
N
Hesperilla idothea clara
Orange grass
Rare
Generalist
Y
Taractrocera anisomorpha
dart
White banded
Rare
Generalist
Y
Taractrocera papyria papyria
grass dart
Eliena rush
Vulnerable
Generalist
N
Trapezites eliena
Rare White spot Vulnerable
Generalist
N
Trapezites luteus luteus
rush
Phigalia rush
Vulnerable
Generalist
N
Trapezites phigalia
Symmomus rush Vulnerable
Specialist
N
Trapezites symmomus soma
North America
Arogos
Endangered (USA states IL, NJ, NY),
Generalist?
Y
Atrytone arogos
Threatened (USA state MN), Species
of Concern (USA state IA)
Light spotted
Data Deficient, rare (Costa Rica,
Dalla octomaculata
Reference
(3)
(3)
(3)
(6)
(3)
(7)
(4)
(4)
(4)
(4)
(4)
(4)
(4)
(4)
(4)
(4)
(4)
(10, 16)
(6)
41
Species
Common Name
Euphyes dukesi
Hesperia dacotae
Dukes’
Dakota
Hesperia leonardus montana
Hesperia ottoe
Pawnee
montane
Otto
Oarisma poweshiek
Poweshiek ling
Panoquina errans
Polites mardon
Wandering
Mardon
Problema bulenta
Problema byssus
Pseudocopaeodes eunus
obscurus
Pyrgus ruralis lagunae
Rare
Byssus
Carson
wandering
Laguna
mountains
Taiwan/Japan
Caltoris bromus yanuca
Fruhstorfer
Carterocephalus palaemon
palaemon
Chequered
Status
Panama)
Threatened (US state (MI)
Threatened (Canada), Endangered
(Canada province MB), Federal
Candidate (USA), Vulnerable (ICUN)
Threatened (USA Federal listing)
Larval Diet
Hesperiinae?
Reference
Generalist
Generalist
Y
Y
(15, 16)
(6, 14)
Generalist
Y
(6, 12)
Endangered (Canada), Threatened
(Canada province MB), Threatened
(USA states (IL, MI, MN)
Threatened (Canada), Endangered
(USA state WI), Threatened (USA
state IA, MI), Species of Concern
(USA state MN)
Threatened (Mexico, USA)
Endangered (USA state WA &
Federal Candidate USA)
Endangered (US state DE)
Threatened (US state IA)
Endangered (USA Federal listing)
Generalist
Y
(10, 16)
Generalist
Y
(2, 13, 16)
Specialist
Generalist
Y
Y
(6)
(1, 8)
?
?
Specialist
Y
Y
Y
(10, 16)
(10, 16)
(13, 16)
-
N
(1, 11)
Extremely Rare, Taiwan
Specialist
-
(5)
Priority (Scotland), Endangered
(Japan)
Specialist
N
(3)
Endangered (USA Federal listing)
Appendix A References
1. Black, S. H., and D. M. Vaughan. 2005. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds). Red List of
Pollinator Insects of North America. CD-ROM Version 1 (May 2005). Portland, OR: The Xerces Society for
Invertebrate Conservation.
2. COSEWIC. 2003. COSEWIC assessment and status report on the Powesheik skipperling Oarisma poweshiek in
Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. vi + 25 pp.
3. Eeles, P. 2002. "U.K. Butterflies". Last updated 2008. Date of access November 28, 2008.
http://www.ukbutterflies.co.uk/species.php?
4. GRUND R., 1998. “South Australian Butterflies”. Last update 4 October 2004. Date of access November 21, 2008.
http://users.sa.chariot.net.au/~rgrund/checklist.htm
42
5. Hsu, Y. F., Wang, S. M. 2004. Notes on immature biology of a rare skipper Caltoris bromus yanuca Fruhstorfer
(Lepidoptera : Hesperiidae), with a discussion on the systematic status of Parnara ranrunna Sonan. Pan-Pacific
Entomologist. 80: 69-80.
6. ICUN 2008. "International Union for Conservation of Nature and Natural Resources" Date of access December 1,
2008. http://www.iucnredlist.org/
7. Opler et al 2008. "The butterflies and skippers of North America". Date of access December 1, 2008.
http://www.nearctica.com/butter/bylist/lhesp.htm
8. Potter, A., J. Fleckenstein, S. Richardson, and D. Hays. 1999. Washington State Status Report for the Mardon
Skipper. Washington Department of Fish and Wildlife, Olympia 39 pp.
9. Shepard, J.H. 2000. Status of five butterflies and skippers in British Columbia. B.C. Minist. Environ., Lands and Parks,
Wildl. Branch and Resour. Inv. Branch, Victoria, BC. 27pp.
10. Shepherd, M. D. 2005. In Shepherd, M. D., D. M. Vaughan, and S. H. Black (Eds). Red List of Pollinator Insects of
North America. CD-ROM Version 1 (May 2005). Portland, OR: The Xerces Society for Invertebrate Conservation.
11. USFWS. 1997. Endangered and threatened wildlife and plants; determination of endangered status for the Laguna
Mountains skipper and Quino Checkerspot Butterfly. Federal Register.62,2313-2322
12. USFWS. 1998. Pawnee montane skipper butterfly (Hesperia leonoardus montana) recovery plan. Denver, Colorado. .
Page 16 pp.
13. USFWS. 2005. Draft recovery plan for the Carson wandering skipper (Pseudocopaedes eunus obscurus).U.S. Fish
and Wildlife Service, Portland, Oregon. viii:71 pp.
14. USFWS 2008."Endangered Species Program" Last updated December 3, 2008. Date of access December 7, 2008.
http://www.fws.gov/northdakotafieldoffice/endspecies/species/dakota_skipper.htm
43
15. Vaughan, D. M., and M. D. Shepherd. 2005. Species Profile: Euphyes dukesi. In Shepherd, M. D., D. M. Vaughan, and
S. H. Black (Eds). Red List of Pollinator Insects of North America. CD-ROM Version 1 (May 2005). Portland, OR:
The Xerces Society for Invertebrate Conservation.
16. Xerces 2008. "The Xerces Society for Invertebrate Conservation; Red List of Butterflies and moths". Date of access
December 7, 2008. http://www.xerces.org/red-list-of-butterflies-and-moths/
Appendix B: Measured Habitat Variables
Scale
Oviposition Plant
Variable
Maximum Culm height (cm)
Basal leaf height (cm)
Percent brown or dead
Foot print area (cm2)
Distance to nearest neighboring plant of the same species (cm)
Structure (Category 1-solitary, 2-loose, 3-dense, 4-matted)
Quadrat (1m2)
Percent cover of bare ground
Percent cover over story canopy
Percent cover of cryptograms
Percent cover of graminoids combined
Percent cover of each species of graminoid
Percent cover of forbs combined
Percent cover of litter
Percent cover of rocks/pebbles
Horizontal Vegetation thickness (0.3, 0.6, & 0.9 meter increments)
Maximum plant height (cm)
Maximum height of each species of graminoid (cm)
Number of blooming forbs (potential nectar resources)
Graminoid species richness (Count of species present)
Graminoid Heterogeneity (Shannon-diversity calculation)
Graminoid Evenness (Shannon-diversity calculation)
Sub-plot (1/10th m2)
Soil Ph
Soil Moisture retention capacity
Litter Depth (cm)
Percent cover of graminoids combined
Percent cover of each species of graminoid
Percent cover of forbs combined
Maximum height of each species of graminoid (cm)
Graminoid species richness (Count of species present)
Graminoid Heterogeneity (Shannon-diversity calculation)
Graminoid Evenness (Shannon-diversity calculation)
Landscape
Number of blooming forbs (potential nectar resources) sampled from 5 m radius
Distance to nearest visible water source (m)
Distance to nearest tree (m)
Distance to nearest shrub (m)
Sapling abundance within 10m of quadrat
Seedling abundance within 10m of quadrat
Tree abundance within 10m of quadrat
Distance to forest edge (m)
Percent slope with regards to 20 m radius of quadrat
44
Appendix C: 2007 Population Counts
Meadow
Date
Count
7A
8-Jul
49
25-Jul
32
1-Aug
7
Cave Creek 31-May
31
8-Jun
56
18-Jun
39
23-Jun
0
Flog Salvage 17-Jun
23
22-Jun
23
Grapefern
9-Jul
313
26-Jul
37
1-Aug
2
Lost
12-Jun
15
18-Jun
14
26-Jun
9
Midway
8-Jul
54
25-Jul
5
1-Aug
0
Muddy
12-Jul
91
26-Jul
31
2-Aug
10
Peterson
8-Jun
23
18-Jun
34
23-Jun
17
Smith Butte
1-Jul
15
3-Jul
37
45
Appendix D: Multiple Consecutive Ovipositions
Meadow
7A
Cave Creek
Flog Salvage
46
Grapefern
Lost
Midway
Muddy
Peterson
Smith Butte
Individual
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
Oviposition Plant Species
Carex fracta, C. fracta/Dantonia intermedia mix
Carex deflex, C. fracta
Danthonia intermedia, Muhlenbergia filiformis
Festuca rubra
F. rubra
Poa pratensis, P. pratensis/F. rubra mix
Carex hoodii, P. pratensis
F.rubra
Festuca roemeri
F.roemeri
F. roemeri
Carex inops
C. deflex, C.deflexa/P. pratensis mix
C. deflexa, P. pratensis
P. pratensis
D. intermedia, C. deflexa
C.deflexa, P. pratensis
P. pratensis, P. pratensis/Bromus carinatus mix
D. intermedia, D. intermedia/Deschampsia cespitosa mix
Carex hallinana, P. pratensis
F. roemeri
F. roemeri
F. roemeri
No. Consecutive
Ovipositions
2
2
2
2
3
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
3
3
2
5
Appendix E: Within Meadow DFA Variable Descriptions
DFA Total Structure Coefficients, level of significance, mean and standard deviations for
each variable in oviposition and random locations, and unit of measurement.
7A Meadow
Variable
Tree abundance within 10 m of quadrat
Total graminoid cover (Egg vicinity)
Distance to nearest visible water source
Oviposition plant leaf density
Canopy cover
Poa pratensis cover (Quadrat)
Distance to nearest tree
Percent slope relative to 20 m radius of quadrat
Litter depth (Egg vicinity)
Carex fracta cover (Egg vicinity)
Oviposition plant footprint
Carex fracta height (Quadrat)
Carex fracta cover (Quadrat)
Graminoid species richness (Egg vicinity)
Poa pratensis height (Quadrat)
Muhlenbergia filiformis height (Egg vicinity)
Litter cover (Quadrat)
Oviposition plant nearest neighbor
Danthonia intermedia height (Egg vicinity)
Poa pratensis cover (Egg vicinity)
Number of blooming forbs within 5 m radius
Total graminoid cover (Quadrat)
Oviposition plant maximum culm height
Oviposition plant dead basal leaves
Bare ground cover (Quadrat)
Muhlenbergia filiformis cover (Egg vicinity)
Carex fracta height (Egg vicinity)
Soil Ph (Egg vicinity)
Stipa occidentalis cover (Egg vicinity)
Danthonia intermedia cover (Egg vicinity)
Cryptogram cover (Quadrat)
Soil moisture (Egg vicinity)
Graminoid heterogeneity (Egg vicinity)
Muhlenbergia filiformis cover (Quadrat)
Agrostis thurberiana cover (Quadrat)
Shrub abundance within 10 m
Rock/pebble cover (Quadrat)
Danthonia intermedia cover (Quadrat)
Stipa occidentalis cover (Quadrat)
Forb cover (Quadrat)
Number of blooming forbs (Quadrat)
Poa pratensis height (Egg vicinity)
Stipa occidentalis height (Quadrat)
Stipa occidentalis height (Egg vicinity)
Agrostis thurberiana cover (Egg vicinity)
Horizontal vegetation thickness at 0.3 m height
Maximum plant height (Quadrat)
Agrostis thurberiana height (Egg vicinity)
Danthonia intermedia height (Quadrat)
Agrostis thurberiana height (Quadrat)
Graminoid species richness (Quadrat)
Oviposition plant basal leaf height
Graminoid species evenness (Egg vicinity)
Muhlenbergia filiformis height (Quadrat)
Graminoid species evenness (Quadrat)
Forb cover (Egg vicinity)
Distance to forest edge
Graminoid heterogeneity (Quadrat)
Vasscular plant cover (Quadrat)
Sapling abundance within 10 m of quadrat
Seedling abundance within 10 m of quadrat
Location
Total Structure
Coefficient Significance Oviposition
Random
-0.486
<0.0001
8.5 ± 7.3
18.0 ± 10.0
0.460
<0.0001
34.8 ± 24.6
20.5 ± 27.1
0.440
<0.01
89.1 ± 0.9
64.9 ± 1.1
0.439
<0.01
2.2 ± 0.3
1.5 ± 0.2
-0.433
<0.05
5.5 ± 109.9
13.3 ± 93.2
0.315
<0.05
5.0 ± 3.3
3.2 ± 2.9
0.312
<0.05
6.9 ± 2.7
5.2 ± 2.8
0.312
<0.05
11.0 ± 6.0
8.3 ± 6.7
0.307
<0.05
1.6 ± 0.7
1.2 ± 0.8
0.280
6.8 ± 12.7
1.2 ± 3.9
0.277
<0.05
78.9 ± 11.3
43.3 ± 8.4
0.272
20.3 ± 12.1
13.3 ± 13.3
0.263
<0.05
5.9 ± 6.1
3.3 ± 4.6
0.255
3.6 ± 1.2
3.0 ± 0.9
0.248
25.4 ± 13.5
18.5 ± 14.2
0.247
3.0 ± 11.9
0.2 ± 1.3
0.246
<0.05
68.3 ± 10.3
55.4 ± 7.7
-0.241
<0.05
6.5 ± 20.2
8.2 ± 14.5
0.236
<0.05
17.2 ± 9.5
12.5 ± 10.4
0.229
3.5 ± 3.3
2.0 ± 2.6
0.223
14.2 ± 7.6
11.1 ± 6.3
0.221
<0.05
37.8 ± 4.6
31.5 ± 11.4
0.220
6.7 ± 0.7
3.0 ± 0.6
0.220
2.8 ± 4.0
1.4 ± 2.8
-0.214
14.2 ± 21.8
20.9 ± 20.0
0.210
3.4 ± 9.7
0.7 ± 3.7
0.208
6.5 ± 11.1
2.0 ± 5.2
-0.188
6.5 ± 9.1
6.6 ± 7.4
0.185
1.6 ± 4.6
0.3 ± 1.0
0.179
11.2 ± 8.5
0.7 ± 3.7
-0.177
12.1 ± 17.0
14.2 ± 18.0
0.171
33.3 ± 19.6
26.1 ± 30.0
0.158
0.4 ± 0.1
0.4 ± 0.1
0.146
4.9 ± 11.9
2.0 ± 4.8
0.142
4.1 ± 7.0
2.1 ± 3.6
0.124
8.3 ± 10.4
5.3 ± 6.2
0.115
0.6 ± 16.9
0.5 ± 20.2
0.114
15.5 ± 10.2
14.1 ± 14.9
0.111
0.8 ± 1.8
0.5 ± 1.3
0.108
37.2 ± 4.6
39.0 ± 3.6
0.108
5.2 ± 4.1
4.8 ± 5.5
0.098
12.2 ± 10.5
10.3 ± 9.2
0.097
5.1 ± 11.2
3.2 ± 9.3
0.091
2.5 ± 6.4
1.9 ± 7.4
0.090
4.0 ± 8.0
2.8 ± 7.6
-0.076
1.5 ± 1.4
1.9 ± 2.6
-0.069
40.3 ± 9.5
42.3 ± 11.9
0.052
20.3 ± 3.9
19.3 ± 7.5
-0.041
30.6 ± 10.0
31.4 ± 11.7
0.037
11.6 ± 14.1
10.7 ± 12.9
0.032
6.3 ± 1.7
6.2 ± 1.4
-0.032
11.7 ± 11.5
12.1 ± 8.7
-0.031
0.8 ± 0.2
0.8 ± 0.2
0.029
1.8 ± 3.0
2.6 ± 5.8
0.017
0.8 ± 0.1
0.8 ± 0.1
0.016
40.8 ± 23.2
40.2 ± 19.9
0.008
17.5 ± 14.0
16.9 ± 16.2
0.007
0.7 ± 0.1
0.7 ± 0.1
64.5 ± 17.4
59.7 ± 15.5
0.0 ± 0.0
0.0 ± 0.0
0.0 ± 0.0
0.1 ± 0.3
3.1 ± 4.0
4.4 ± 3.7
2.6 ± 2.7
5.7 ± 4.5
47
Unit
count
%
meters
category (1-4)
%
%
meters
%
cm
%
2
cm
cm
%
count
cm
cm
%
cm
cm
%
count
%
cm
%
%
%
cm
scale
%
%
%
%
index
%
%
count
%
%
%
%
count
cm
cm
cm
%
%
cm
cm
cm
cm
count
cm
index
cm
index
%
meters
index
%
%
%
count
count
Cave Creek Meadow
Variable
Festuca rubra cover (Egg vicinity)
Canopy cover
Festuca rubra cover (Quadrat)
Carex hoodii height (Quadrat)
Carex hoodii cover (Quadrat)
Festuca rubra height (Egg vicinity)
Carex hoodii cover (Egg vicinity)
Carex hoodii height (Egg vicinity)
Festuca rubra height (Quadrat)
Location
Total Structure
Coefficient
0.656944
-0.588236
0.539355
-0.473703
0.455402
-0.452389
0.407111
-0.359243
0.32858
0.325884
-0.315773
0.312784
-0.311138
0.308497
0.287832
-0.279337
-0.271652
-0.260067
0.258576
0.254345
-0.253685
-0.238895
0.233471
-0.206937
0.206375
0.206001
0.201905
0.197964
-0.187409
-0.169455
-0.166846
0.16112
0.154348
-0.153629
0.146338
0.121675
-0.094334
-0.087642
-0.087043
0.071653
0.064973
-0.064289
0.060235
0.052678
0.048411
0.04409
0.011476
-
48
Significance
<0.0001
<0.0001
<0.0001
<0.0001
<0.0001
<0.001
<0.0001
<0.01
<0.05
<0.05
<0.05
<0.05
<0.05
<0.05
<0.05
<0.05
<0.05
<0.05
<0.01
<0.05
-
Oviposition
38.4 ± 6.2
2.7 ± 0.4
1.5 ± 1.6
28.9 ± 22.0
5.4 ± 80.3
9.2 ± 4.8
10.8 ± 16.9
47.0 ± 35.6
0.4 ± 0.8
0.1 ± 0.4
14.3 ± 15.8
3.1 ± 0.5
21.4 ± 17.5
72.8 ± 17.1
30.5 ± 11.8
25.6 ± 14.9
85.1 ± 11.5
0.6 ± 10.4
9.6 ± 17.3
80.5 ± 1.1
11.7 ± 11.0
46.4 ± 23.2
0.7 ± 0.3
17.6 ± 10.3
0.3 ± 0.1
34.5 ± 2.1
52.6 ± 16.7
0.2 ± 0.9
12.5 ± 0.7
14.0 ± 14.4
7.5 ± 17.7
1.1 ± 5.4
20.4 ± 9.3
17.0 ± 21.3
27.9 ± 22.0
20.5 ± 20.2
68.9 ± 24.9
3.7 ± 3.3
15.1 ± 10.7
6.2 ± 9.7
1.4 ± 0.7
5.2 ± 10.7
50.0 ± 7.4
6.8 ± 22.3
6.7 ± 8.7
23.6 ± 11.4
2.6 ± 0.6
0.8 ± 0.1
0.4 ± 0.1
0.0 ± 0.0
0.0 ± 0.0
Random
52.8 ± 10.8
1.8 ± 0.3
4.7 ± 4.5
10.9 ± 16.7
16.4 ± 68.9
5.1 ± 3.3
18.7 ± 3.4
29.5 ± 28.9
4.3 ± 10.1
0.7 ± 1.3
9.4 ± 14.4
3.5 ± 0.9
12.9 ± 15.0
81.2 ± 20.9
39.1 ± 17.6
15.9 ± 19.7
47.8 ± 16.5
0.1 ± 11.1
26.6 ± 35.0
94.0 ± 0.5
8.3 ± 13.5
36.7 ± 28.6
0.8 ± 0.3
12.7 ± 13.5
0.3 ± 0.1
43.0 ± 2.5
62.1 ± 8.5
1.6 ± 7.3
10.8 ± 0.8
10.6 ± 12.7
3.1 ± 19.6
1.8 ± 8.1
24.0 ± 14.6
9.6 ± 9.8
33.7 ± 22.6
25.1 ± 22.8
65.3 ± 18.9
3.3 ± 4.3
14.7 ± 13.9
6.2 ± 14.0
2.1 ± 3.4
4.9 ± 10.3
52.3 ± 17.1
4.4 ± 18.1
7.7 ± 13.9
25.0 ± 20.7
2.6 ± 0.7
0.8 ± 0.1
0.4 ± 0.1
1.0 ± 5.5
0.1 ± 0.6
Unit
cm
category (1-4)
count
%
%
meters
cm
count
count
count
%
index
%
%
cm
cm
2
cm
%
%
meters
%
%
index
cm
index
%
%
%
cm
%
%
%
cm
count
%
meters
%
%
%
scale
cm
%
%
cm
cm
cm
index
index
index
%
count
Flog Salvage meadow
Variable
Festuca idahoensis cover (Egg vicinity)
Festuca idahoensis height (Egg vicinity)
Festuca idahoensis cover (Quadrat)
Festuca idahoensis height (Quadrat)
Carex inops cover (Quadrat)
Carex inops height (Quadrat)
Canopy cover
Carex inops cover (Egg vicinity)
Carex inops height (Egg vicinity)
Location
Total Structure
Coefficient Significance
0.783
<0.0001
0.772
0.709
0.679
0.614
0.609
0.591
0.568
0.565
0.480
0.453
0.437
0.435
0.434
0.430
0.385
0.378
-0.301
-0.298
0.294
0.275
0.262
-0.261
-0.244
-0.218
-0.194
-0.182
-0.180
0.162
-0.155
0.142
0.115
0.112
-0.110
0.105
0.101
0.097
-0.062
0.054
-0.052
0.035
0.032
0.030
-0.019
-0.015
0.012
<0.0001
<0.0001
<0.0001
<0.0001
<0.0001
<0.0001
<0.0001
<0.0001
<0.01
<0.01
<0.0001
<0.01
<0.01
<0.01
<0.01
<0.01
<0.05
<0.05
<0.05
-
49
Oviposition
30.8 ± 22.1
212.3
2.8
42.3
34.5
2.3
12.8
18.8
11.0
12.3
0.2
20.7
3.0
0.6
7.6
0.4
44.3
85.3
56.2
0.7
19.6
13.5
0.8
17.4
0.5
3.5
2.4
31.6
21.5
67.6
7.9
0.9
21.2
6.4
1.9
15.3
3.2
11.3
6.8
12.2
0.1
0.7
2.6
14.2
43.1
3.0
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
9.8
0.2
24.0
6.4
0.6
7.0
20.9
6.5
0.4
0.1
13.8
0.7
0.3
6.3
0.1
7.0
31.3
22.4
0.2
12.4
19.8
3.0
4.5
2.7
2.7
6.2
12.9
12.5
3.2
8.2
14.6
20.9
168.1
11.8
13.2
1.4
7.2
30.4
11.6
4.2
1.2
0.5
7.4
19.8
3.6
Random
2.0 ± 6.5
30.7
1.6
16.5
20.3
1.4
2.8
3.9
3.7
6.7
0.1
8.5
2.2
0.3
3.1
0.2
33.5
105.8
71.7
0.6
13.0
11.6
11.1
20.3
7.9
6.2
9.6
55.9
18.3
75.4
5.1
0.7
17.2
8.4
1.7
12.6
3.3
12.5
6.8
12.4
0.1
0.9
1.4
14.4
43.4
2.7
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
5.9
0.2
28.6
9.4
0.6
7.9
12.5
6.0
0.3
0.1
13.1
1.0
0.4
4.5
0.2
4.7
37.3
29.6
0.4
8.0
3.5
26.6
7.1
24.6
9.1
21.3
71.5
7.3
5.1
3.8
17.5
12.5
29.1
8.0
15.7
2.8
9.6
28.7
8.6
4.5
2.0
0.7
5.3
10.6
3.0
Unit
%
cm2
category (1-4)
%
%
count
cm
cm
%
%
index
cm
count
index
%
index
%
count
count
index
%
cm
%
cm
%
%
%
cm
count
%
meters
%
%
%
%
%
%
count
scale
%
%
count
cm
cm
meters
count
Grapefern Meadow
Variable
Carex inops cover (Quadrat)
Canopy cover
Carex inops height (Egg vicinity)
Carex inops height (Quadrat)
Carex inops cover (Egg vicinity)
Bromus carinatus cover (Quadrat)
Bromus carinatus cover (Egg vicinity)
Bromus carinatus height (Quadrat)
Elymus elymoides cover (Egg vicinity)
Elymus elymoides height (Egg vicinity)
Bromus carinatus height (Egg vicinity)
Elymus elymoides cover (Quadrat)
Elymus elymoides height (Quadrat)
Location
Total Structure
Coefficient
Significance Oviposition
Random
Unit
0.667
<0.0001
2.1 ± 0.7
0.8 ± 0.9
cm
0.605
<0.0001
48.8 ± 15.5 27.0 ± 12.8
%
0.565
<0.0001
82.5 ± 28.0 53.8 ± 10.9
%
0.538
<0.0001
50.8 ± 19.2 27.5 ± 17.1
%
-0.484
<0.0001
3.0 ± 20.9 16.3 ± 3.9
count
-0.476
<0.0001
2.3 ± 10.7
9.5 ± 3.3
count
0.467
<0.0001
65.8 ± 16.3 50.2 ± 13.1
%
0.452
<0.01
20.5 ± 15.6
6.9 ± 12.1
%
-0.435
<0.01
6.9 ± 24.4 25.8 ± 6.7
%
0.412
27.5 ± 9.5
18.5 ± 10.7
meters
-0.386
<0.05
1.0 ± 10.6
7.2 ± 1.3
%
0.367
<0.01
17.4 ± 5.7
12.7 ± 6.3
cm
0.365
<0.01
13.7 ± 10.2
6.5 ± 8.5
cm
0.352
<0.01
2.2 ± 0.8
1.7 ± 0.8
category (1-4)
0.343
<0.01
19.3 ± 12.7 10.9 ± 10.6
cm
-0.342
<0.05
7.4 ± 31.2 23.2 ± 17.2
count
0.335
<0.01
23.8 ± 20.3 10.2 ± 18.8
%
-0.321
<0.01
6.4 ± 0.3
6.6 ± 0.3
Scale
0.316
<0.01
15.1 ± 6.6
11.3 ± 4.6
meters
0.303
<0.05
22.3 ± 17.7 11.8 ± 15.8
cm
0.278
<0.05
6.7 ± 5.9
3.9 ± 8.1
%
-0.255
<0.05
0.6 ± 11.8
4.9 ± 1.6
count
0.249
5.1 ± 4.6
3.0 ± 3.6
%
0.245
<0.05
108.2 ± 59.2 46.3 ± 186.5
cm2
0.236
<0.05
8.1 ± 11.6
3.3 ± 8.4
%
0.233
7.8 ± 4.3
5.6 ± 6.1
count
-0.225
8.5 ± 10.6 12.7 ± 7.2
%
0.218
0.3 ± 0.2
0.2 ± 0.1
Index
0.213
2.8 ± 1.1
2.4 ± 0.7
Index
-0.209
2.0 ± 7.1
3.6 ± 6.6
cm
0.205
26.7 ± 8.0
23.0 ± 9.9
%
0.202
0.7 ± 0.4
0.6 ± 0.2
Index
0.179
4.0 ± 4.3
2.9 ± 4.0
%
0.169
10.1 ± 12.3
6.4 ± 7.6
%
0.153
6.5 ± 15.1
2.3 ± 6.5
%
0.147
19.2 ± 14.8 14.8 ± 15.3
cm
0.137
45.7 ± 15.9 44.3 ± 8.7
cm
-0.132
0.8 ± 0.1
0.8 ± 0.1
Index
-0.127
1.3 ± 4.3
2.0 ± 4.0
%
0.121
4.6 ± 1.5
4.3 ± 1.3
Index
0.118
3.7 ± 13.2
1.4 ± 4.9
%
-0.117
2.9 ± 4.1
4.1 ± 2.9
%
0.114
29.4 ± 19.4 29.0 ± 26.4
cm
0.113
5.2 ± 13.5
2.8 ± 7.3
cm
0.100
1.5 ± 5.8
0.5 ± 2.0
%
0.086
28.0 ± 17.6 26.5 ± 12.2
%
0.086
9.1 ± 11.9
7.2 ± 10.4
cm
0.085
2.3 ± 2.7
1.9 ± 3.3
count
0.073
5.8 ± 11.0
4.0 ± 9.3
cm
-0.070
4.8 ± 4.4
5.5 ± 4.2
cm
-0.069
3.2 ± 6.6
3.3 ± 4.5
%
-0.068
6.3 ± 2.4
6.7 ± 2.6
%
0.056
2.8 ± 6.4
1.8 ± 3.7
%
0.054
0.5 ± 0.2
0.5 ± 0.1
Index
0.053
3.6 ± 6.9
3.0 ± 5.1
%
-0.046
10.4 ± 15.2 11.7 ± 13.7
cm
0.030
2.1 ± 7.0
1.7 ± 6.5
cm
0.024
23.1 ± 15.9 24.0 ± 10.3
%
-0.022
5.4 ± 14.4
6.5 ± 11.9
cm
0.009
12.0 ± 22.7 11.6 ± 21.3
cm
0.009
12.7 ± 12.1
9.3 ± 17.5
cm
-0.008
2.2 ± 4.4
2.3 ± 5.7
%
0.002
9.8 ± 16.2
9.7 ± 15.0
cm
0.1 ± 1.0
0.3 ± 0.4
%
0.0 ± 0.0
0.0 ± 0.0
%
0.0 ± 0.3
0.1 ± 0.0
%
1.83 ± 10
0 ± 0
%
0.67 ± 3.7
0 ± 0
%
0.67 ± 3.7
0 ± 0
cm
1.33 ± 7.3
0 ± 0
cm
50
Location
Lost Meadow
Variable
Danthonia unispicata cover (Quadrat)
Carex deflexa cover (Egg vicinity)
Danthonia unispicata cover (Egg vicinity)
Danthonia unispicata height (Quadrat)
Carex deflexa cover (Quadrat)
Danthonia unispicata height (Egg vicinity)
Carex deflexa height (Egg vicinity)
Canopy cover
Carex deflexa height (Quadrat)
Total Structure
Coefficient
-0.592
-0.540
-0.479
-0.476
0.471
-0.470
-0.448
-0.442
-0.410
-0.366
-0.355
-0.352
-0.341
-0.333
0.333
-0.329
0.325
-0.319
-0.311
0.298
-0.285
-0.281
-0.280
-0.278
-0.274
-0.273
-0.234
-0.211
0.195
0.194
0.189
-0.177
0.142
-0.142
-0.137
0.130
-0.124
-0.124
-0.108
-0.096
-0.091
-0.088
-0.084
0.067
-0.042
-0.041
-0.038
-0.025
-0.023
-0.021
-
51
Significance
<0.0001
<0.0001
<0.0001
<0.0001
<0.01
0.01
<0.01
<0.01
<0.01
<0.01
<0.05
<0.05
<0.01
<0.05
<0.05
<0.05
<0.05
<0.05
<0.01
<0.05
-
Oviposition
54.7 ± 18.2
53.2 ± 18.3
68.5 ± 16.7
15.9 ± 9.9
10.3 ± 6.5
3.0 ± 0.0
65.1 ± 59.7
2.0 ± 0.0
0.2 ± 0.0
21.5 ± 20.6
12.5 ± 11.4
0.7 ± 0.3
4.2 ± 6.7
11.5 ± 12.3
59.7 ± 11.5
2.3 ± 0.6
27.0 ± 21.4
0.4 ± 0.0
0.5 ± 0.0
0.2 ± 0.4
19.4 ± 22.8
13.1 ± 5.7
6.3 ± 0.4
12.0 ± 12.0
7.4 ± 5.7
6.1 ± 7.2
12.0 ± 6.2
4.1 ± 5.9
7.5 ± 7.4
22.0 ± 8.2
0.6 ± 2.1
7.7 ± 3.2
2.5 ± 2.8
17.2 ± 11.6
9.4 ± 7.5
17.4 ± 7.8
20.1 ± 9.8
16.4 ± 7.0
0.9 ± 1.6
7.9 ± 15.1
26.0 ± 5.5
20.5 ± 6.9
5.9 ± 4.8
2.8 ± 1.6
42.5 ± 23.7
8.8 ± 8.9
7.9 ± 10.3
0.9 ± 0.0
6.9 ± 11.4
1.1 ± 2.2
0.1 ± 0.2
0.0 ± 0.0
0.0 ± 0.0
0.2 ± 0.8
0.0 ± 0.0
1.9 ± 5.9
1.7 ± 4.8
7.2 ± 9.9
Random
Unit
29.1 ± 19.2
%
32.3 ± 16.3
%
51.3 ± 16.9
%
8.1 ± 5.5
%
28.3 ± 22.9
%
3.0 ± 0.0
count
2
cm
28.5 ± 42.0
2.0 ± 0.0
count
0.2 ± 0.0
index
10.3 ± 14.9
%
6.3 ± 6.0
%
0.5 ± 0.5
cm2
0.7 ± 2.2
%
4.3 ± 6.6
%
68.6 ± 13.8
%
1.9 ± 0.6 category (1-4)
42.6 ± 29.4
count
0.4 ± 0.0
index
0.5 ± 0.0
index
1.1 ± 2.2
%
9.2 ± 14.8
%
8.9 ± 9.0
cm
6.1 ± 0.4
scale
6.0 ± 6.0
%
3.5 ± 5.1
cm
3.0 ± 4.2
cm
9.1 ± 6.6
cm
2.3 ± 5.4
cm
10.4 ± 11.0
count
25.7 ± 10.6
%
1.7 ± 3.5
count
7.1 ± 4.5
cm
4.4 ± 9.2
%
14.6 ± 12.0
%
7.6 ± 6.0
%
19.6 ± 8.1
meters
17.5 ± 11.4
cm
14.7 ± 7.0
meters
0.7 ± 0.9
%
6.1 ± 7.6
%
25.1 ± 6.3
cm
19.1 ± 9.5
cm
5.3 ± 4.9
cm
3.9 ± 5.4
cm
40.9 ± 29.3
%
9.9 ± 15.6
%
7.2 ± 8.9
cm
0.9 ± 0.0
index
5.8 ± 8.5
%
1.2 ± 2.8
count
0.1 ± 0.3
%
0.0 ± 0.0
%
0.0 ± 0.0
%
0.3 ± 1.3
count
0.0 ± 0.0
count
1.0 ± 3.8
%
0.7 ± 2.9
cm
2.2 ± 5.9
cm
Muddy Meadow
Variable
Agrostis thurberiana cover (Quadrat)
Agrostis thurberiana height (Quadrat)
Muhlenbergia filiformis cover (Quadrat)
Agrostis thurberiana cover (Egg vicinity)
Muhlenbergia filiformis height (Quadrat)
Muhlenbergia filiformis cover (Egg vicinity)
Calamagrostis canadensis height (Quadrat)
Agrostis thurberiana height (Egg vicinity)
Calamagrostis canadensis height (Egg vicinity)
Canopy cover
Muhlenbergia filiformis height (Egg vicinity)
Calamagrostis canadensis cover (Egg vicinity)
Calamagrostis canadensis cover (Quadrat)
Location
Total Structure
Coefficient
Random
Unit
-0.477
<0.0001
6.1 ± 0.3
6.4 ± 0.3
Scale
0.474
<0.0001
6.4 ± 1.4
4.9 ± 1.5
Index
0.467
<0.01
29.9 ± 13.6
13.6 ± 17.7
cm
0.441
<0.0001
20.5 ± 13.7
9.2 ± 17.3
%
0.429
<0.01
17.1 ± 24.7
3.5 ± 6.2
%
0.422
<0.01
92.4 ± 12.9
79.8 ± 17.5
%
0.409
<0.01
17.4 ± 12.0
6.8 ± 12.1
cm
0.407
<0.01
59.7 ± 19.1
41.0 ± 23.8
%
-0.397
<0.01
4.2 ± 4.6
15.0 ± 17.6
%
0.374
<0.01
12.3 ± 13.6
5.1 ± 13.5
%
0.368
<0.01
3.9 ± 1.1
3.0 ± 1.0
index
0.358
<0.01
19.2 ± 12.1
9.0 ± 14.9
cm
0.349
<0.05
20.2 ± 22.6
7.5 ± 12.2
%
0.348
<0.05
56.7 ± 20.7
39.6 ± 27.6
%
-0.347
<0.05
3.2 ± 6.7
12.0 ± 15.3
%
0.323
<0.05
51.4 ± 11.5
42.3 ± 15.2
%
-0.319
<0.05
0.1 ± 0.3
0.7 ± 1.4
%
0.314
<0.01
0.6 ± 0.1
0.5 ± 0.1
index
0.304
<0.05
10.8 ± 14.5
3.8 ± 10.6
cm
-0.300
<0.05
11.8 ± 22.1
31.1 ± 33.1
cm
-0.295
<0.05
11.4 ± 6.4
16.4 ± 9.1
cm
-0.280
<0.05
4.8 ± 10.9
15.8 ± 22.2
cm
-0.279
<0.05
47.3 ± 11.1
56.5 ± 16.7
cm
-0.277
1.0 ± 2.2
3.1 ± 4.2
%
0.255
12.9 ± 26.6
2.3 ± 4.9
%
-0.220
<0.05
4.0 ± 6.6
6.1 ± 7.9
cm
0.213
2.3 ± 0.9
1.9 ± 0.8
category (1-4)
0.204
5.5 ± 4.7
3.5 ± 5.1
cm
0.202
6.0 ± 6.6
3.6 ± 3.7
%
-0.196
29.7 ± 8.7
38.1 ± 17.4
meters
-0.191
6.6 ± 19.5
12.5 ± 20.8
%
0.175
28.2 ± 27.8
21.9 ± 26.8
count
0.174
12.8 ± 21.8
6.8 ± 12.7
%
0.167
0.4 ± 0.2
0.4 ± 0.2
index
0.163
59.2 ± 23.8
52.5 ± 19.2
%
0.160
3.9 ± 10.7
1.9 ± 10.4
cm
0.158
14.0 ± 23.5
7.5 ± 10.8
%
0.153
52.8 ± 24.5
46.5 ± 20.4
%
0.139
7.6 ± 10.7
4.8 ± 9.4
cm
-0.135
4.7 ± 8.5
6.7 ± 12.0
count
0.134
11.9 ± 5.8
10.1 ± 4.5
meters
-0.133
2.7 ± 10.2
5.5 ± 15.5
%
0.133
4.3 ± 4.1
3.5 ± 4.8
%
-0.117
1.4 ± 3.3
2.6 ± 5.2
count
0.113
1.0 ± 5.3
1.9 ± 10.4
cm
-0.108
5.3 ± 6.0
6.6 ± 6.8
%
-0.075
1.4 ± 2.4
2.1 ± 4.3
count
-0.074
0.7 ± 0.3
0.7 ± 0.3
index
0.070
3.1 ± 4.7
2.7 ± 4.0
%
-0.059
0.8 ± 0.1
0.8 ± 0.1
index
-0.054
80.2 ± 24.8
83.1 ± 23.2
%
0.053
3.1 ± 3.7
2.6 ± 4.6
cm
-0.037
7.4 ± 14.7
6.9 ± 9.3
count
0.035
24.6 ± 16.5
23.4 ± 19.4
cm
cm2
-0.019
78.6 ± 113.6
87.4 ± 133.5
-0.019
1.8 ± 1.1
1.9 ± 1.3
cm
0.014
36.1 ± 16.6
35.7 ± 16.2
meters
-0.014
9.8 ± 11.0
10.2 ± 15.8
cm
0.0 ± 0.0
0.0 ± 0.0
%
0.1 ± 0.4
0.2 ± 0.7
%
0.3 ± 1.8
0.9 ± 3.8
count
0.8 ± 4.6
0.3 ± 1.8
%
2.0 ± 6.5
1.2 ± 6.4
%
52
Midway Meadow
Variable
Bromus carinatus height (Egg vicinity)
Hordeum brachyantherum cover (Quadrat)
Bromus carinatus cover (Quadrat)
Distance to nearest visible water source (m)
Bromus carinatus cover (Egg vicinity)
Hordeum brachyantherum height (Quadrat)
Bromus carinatus height (Quadrat)
Canopy cover
Hordeum brachyantherum cover (Egg vicinity)
Hordeum brachyantherum height (Egg vicinity)
Location
Total Structure
Coefficient
Random
Unit
-0.456
31.3 ± 9.1
41.1 ± 11.2
cm
0.364
28.8 ± 24.1 15.1 ± 10.8
%
0.354
<0.05
49.0 ± 20.5 36.2 ± 14.6
%
-0.351
<0.05
5.4 ± 6.7
15.8 ± 20.0
%
0.318
<0.05
3.5 ± 3.2
2.3 ± 2.3
count
0.305
23.4 ± 21.7 12.7 ± 11.1
%
0.305
<0.05
32.1 ± 9.6
25.4 ± 12.4
%
0.293
<0.05
2.0 ± 0.6
1.7 ± 0.5 category (1-4)
-0.289
<0.05
39.6 ± 10.6 45.5 ± 11.4
cm
0.287
86.6 ± 11.1 74.9 ± 24.4
%
0.280
1.1 ± 0.7
0.8 ± 0.6
cm
-0.269
0.1 ± 0.3
0.3 ± 0.5
%
0.253
38.8 ± 22.5 29.4 ± 13.8
%
0.232
3.0 ± 8.9
0.2 ± 0.9
%
-0.226
14.3 ± 4.9
20.5 ± 10.6
cm
-0.224
10.6 ± 12.0 16.1 ± 13.2
cm
0.218
70.5 ± 17.3 62.2 ± 20.1
%
-0.199
0.8 ± 0.2
0.8 ± 0.2
Index
0.194
7.5 ± 15.4
2.6 ± 10.1
cm
0.189
33.6 ± 45.4 30.9 ± 61.3
cm2
0.187
3.0 ± 7.5
1.0 ± 4.0
%
-0.184
29.0 ± 16.6 35.7 ± 21.2
%
0.176
12.3 ± 8.9
7.7 ± 6.1
count
-0.173
0.7 ± 0.3
0.8 ± 0.2
Index
0.158
2.9 ± 7.5
0.9 ± 5.1
cm
0.147
10.3 ± 10.7
7.7 ± 8.1
cm
0.136
0.9 ± 2.4
0.4 ± 1.0
count
-0.135
7.3 ± 17.1 10.6 ± 17.5
cm
-0.122
0.5 ± 0.2
0.5 ± 0.2
Index
0.109
4.9 ± 7.8
3.5 ± 5.5
%
-0.099
0.5 ± 1.2
1.1 ± 3.3
%
-0.090
0.4 ± 1.0
0.5 ± 0.9
count
0.087
1.5 ± 1.9
1.6 ± 5.5
%
-0.085
1.6 ± 2.2
2.3 ± 3.9
%
-0.084
11.8 ± 8.5
13.7 ± 10.2
%
0.084
57.0 ± 14.1 55.2 ± 15.4
meters
-0.083
9.5 ± 3.5
10.2 ± 4.9
cm
0.070
27.4 ± 18.0 25.2 ± 18.3
%
-0.069
0.3 ± 0.2
0.4 ± 0.1
Index
0.068
6.1 ± 5.0
5.6 ± 4.4
%
0.062
21.9 ± 10.2 20.7 ± 10.4
meters
-0.060
44.9 ± 21.2 46.7 ± 18.8
meters
0.058
5.1 ± 6.7
4.4 ± 6.0
%
-0.051
1.8 ± 4.0
2.1 ± 4.2
%
-0.044
2.7 ± 7.0
5.3 ± 13.4
cm
0.039
4.3 ± 1.2
4.2 ± 1.5
Index
-0.032
2.8 ± 1.1
2.9 ± 1.1
Index
0.031
3.5 ± 5.3
2.7 ± 2.0
cm
0.028
28.6 ± 15.5 27.7 ± 19.1
cm
-0.021
25.6 ± 12.5 26.1 ± 13.2
cm
-0.019
0.6 ± 1.9
0.6 ± 1.5
%
-0.002
2.1 ± 5.2
3.4 ± 9.0
cm
6.6 ± 0.2
6.6 ± 0.2
Scale
0.0 ± 0.0
0.0 ± 0.0
%
0.0 ± 0.0
0.1 ± 0.3
%
1.5 ± 3.0
1.6 ± 3.6
count
0.0 ± 0.2
0.3 ± 1.8
count
5.0 ± 0.0
5.0 ± 0.0
%
6.4 ± 8.8
11.6 ± 14.1
%
0.3 ± 1.8
0.2 ± 0.9
%
0.3 ± 1.8
0.5 ± 2.9
cm
0.4 ± 1.3
1.9 ± 4.8
%
2.3 ± 7.7
4.0 ± 8.3
cm
53
Peterson Prairie Meadow
Variable
Canopy cover
Carex praticola cover (Quadrat)
Carex praticola height (Egg vicinity)
Carex praticola cover (Egg vicinity)
Carex halliana cover (Egg vicinity)
Carex halliana height (Egg vicinity)
Carex halliana cover (Quadrat)
Carex praticola height (Quadrat)
Carex halliana height (Quadrat)
Danthonia unispicata cover (Egg vicinity)
Danthonia unispicata cover (Quadrate)
Danthonia unispicata height (Egg vicinity)
Danthonia unispicata height (Quadrat)
Location
Total Structure
Coefficient Significance Oviposition
Random
Unit
0.521
<0.0001 54.2 ± 11.8 37.7 ± 17.0
meters
0.509
<0.0001 39.1 ± 12.5 23.5 ± 15.8
meters
-0.501
0.4 ± 1.1
7.3 ± 10.9
%
0.494
<0.0001 17.0 ± 8.8
9.5 ± 5.5
%
0.461
<0.01
36.2 ± 11.9 25.5 ± 10.7
%
0.390
6.8 ± 0.2
6.6 ± 0.3
Scale
0.390
8.2 ± 6.0
4.6 ± 4.3
%
-0.367
<0.01
28.5 ± 7.5
34.1 ± 8.1
cm
0.347
<0.01
24.3 ± 10.8 17.7 ± 10.0
%
0.335
<0.05
7.4 ± 11.7
1.4 ± 5.3
cm
-0.331
<0.05
1.6 ± 3.7
4.8 ± 6.8
%
-0.324
<0.01
0.1 ± 0.3
2.9 ± 6.1
count
0.318
<0.05
82.9 ± 16.3 67.6 ± 27.8
%
0.295
0.8 ± 1.9
0.0 ± 0.2
%
0.279
<0.05
12.3 ± 8.7
7.8 ± 5.8
%
-0.274
28.9 ± 11.8 35.8 ± 16.2
%
-0.273
0.1 ± 0.3
1.4 ± 3.2
count
cm2
0.271
<0.05
50.2 ± 47.7 25.0 ± 35.1
0.265
<0.05
2.5 ± 4.5
0.6 ± 2.8
%
0.257
1.8 ± 0.4
1.5 ± 0.5 category (1-4)
0.255
3.8 ± 8.7
0.8 ± 4.6
cm
-0.248
26.0 ± 12.3 33.0 ± 16.6
%
-0.245
<0.01
0.2 ± 0.9
2.3 ± 8.3
%
-0.225
5.7 ± 8.7
12.0 ± 17.7
%
-0.220
0.0 ± 0.2
0.2 ± 0.6
%
-0.217
31.2 ± 16.4 38.7 ± 19.3
%
-0.213
0.8 ± 0.1
0.9 ± 0.1
Index
0.197
1.6 ± 4.5
0.3 ± 1.8
%
-0.190
9.4 ± 6.9
11.9 ± 7.3
cm
-0.186
<0.05
0.1 ± 0.4
2.4 ± 9.2
count
0.183
3.1 ± 8.5
0.8 ± 4.6
cm
0.181
9.6 ± 8.8
6.9 ± 6.6
%
-0.173
3.1 ± 4.1
6.6 ± 11.6
%
-0.171
12.8 ± 4.2
14.6 ± 5.9
cm
0.162
<0.05
5.5 ± 3.1
4.8 ± 4.7
%
-0.152
0.5 ± 0.1
0.5 ± 0.1
Index
-0.142
23.0 ± 8.7
25.9 ± 12.3
cm
-0.129
5.5 ± 10.1
4.8 ± 3.0
cm
-0.107
0.6 ± 2.0
1.1 ± 2.7
%
-0.092
2.3 ± 5.3
3.3 ± 6.6
cm
0.082
1.5 ± 1.6
1.3 ± 2.1
%
-0.078
2.3 ± 0.6
2.4 ± 0.7
Index
-0.069
14.7 ± 6.4
15.7 ± 8.3
cm
-0.061
4.2 ± 0.8
4.3 ± 0.9
Index
-0.061
21.8 ± 6.4
24.5 ± 10.1
cm
-0.060
0.3 ± 0.1
0.3 ± 0.1
Index
0.043
52.5 ± 35.8 47.5 ± 26.6
count
0.036
8.3 ± 8.0
7.7 ± 8.8
cm
0.028
1.2 ± 0.9
1.2 ± 1.0
cm
0.021
0.8 ± 0.3
0.8 ± 0.3
Index
-0.016
15.5 ± 5.4
15.7 ± 8.0
cm
-0.007
56.5 ± 16.8 56.7 ± 20.7
%
0.002
22.0 ± 22.7 16.8 ± 13.4
count
0.0 ± 0.0
0.0 ± 0.2
%
0.0 ± 0.0
0.0 ± 0.0
%
0.0 ± 0.0
0.6 ± 1.9
count
2.2 ± 2.8
2.9 ± 5.3
%
9.9 ± 9.7
8.4 ± 9.9
cm
0.7 ± 0.9
0.0 ± 0.0
%
0.0 ± 0.0
0.0 ± 0.0
%
0.2 ± 1.3
0.0 ± 0.0
cm
0.0 ± 0.0
0.0 ± 0.0
cm
54
Smith Butte Meadow
Variable
Festuca idahoensis cover (Egg vicinity)
Festuca idahoensis height (Egg vicinity)
Festuca idahoensis cover (Quadrat)
Festuca rubra height (Quadrat)
Festuca rubra cover (Quadrat)
Festuca idahoensis height (Quadrat)
Festuca rubra height (Egg vicinity)
Festuca rubra cover (Egg vicinity)
Canopy cover
Location
Total Structure
Coefficient
Random
-0.694
<0.0001
2.2 ± 0.4
1.5 ± 0.5
-0.686
<0.0001 23.2 ± 14.3
6.5 ± 9.3
-0.649
<0.0001 18.8 ± 7.1
7.9 ± 7.9
-0.505
<0.001
23.5 ± 14.2
11.5 ± 11.2
-0.503
<0.0001 76.4 ± 61.3
40.9 ± 116.0
-0.459
<0.001
16.2 ± 3.9
12.4 ± 4.3
0.418
<0.001
2.1 ± 8.2
13.5 ± 17.8
0.416
<0.01
1.0 ± 4.6
5.9 ± 8.2
-0.404
<0.01
28.0 ± 6.1
22.5 ± 7.7
0.402
<0.05
5.3 ± 4.4
26.6 ± 32.9
0.377
<0.01
0.0 ± 0.0
0.7 ± 1.4
-0.376
<0.01
26.1 ± 11.6
16.4 ± 14.8
0.373
<0.01
0.9 ± 5.1
6.9 ± 11.9
0.364
<0.05
2.5 ± 1.3
4.3 ± 2.8
0.266
46.8 ± 8.4
52.4 ± 12.2
0.263
<0.01
0.3 ± 1.8
3.0 ± 7.4
0.255
7.8 ± 4.8
9.4 ± 3.2
-0.206
6.8 ± 0.1
6.7 ± 0.2
-0.201
12.3 ± 5.1
10.5 ± 4.6
0.182
23.7 ± 7.3
27.4 ± 10.8
0.168
10.4 ± 3.9
12.7 ± 7.0
-0.158
0.4 ± 0.1
0.4 ± 0.1
-0.157
0.6 ± 0.1
0.5 ± 0.1
-0.156
0.8 ± 0.1
0.8 ± 0.2
-0.156
15.7 ± 18.1
12.1 ± 14.1
0.149
53.2 ± 18.1
57.7 ± 16.1
-0.145
4.5 ± 0.6
4.2 ± 1.1
0.127
77.5 ± 22.4
81.5 ± 20.2
-0.118
3.3 ± 0.8
3.1 ± 1.0
0.108
58.0 ± 19.3
61.5 ± 16.1
0.108
0.7 ± 1.3
1.0 ± 1.9
0.085
0.4 ± 0.7
0.6 ± 1.1
-0.081
0.9 ± 0.1
0.8 ± 0.1
0.080
5.9 ± 5.1
6.7 ± 5.4
0.079
16.7 ± 4.7
17.8 ± 9.2
-0.079
0.2 ± 0.6
0.2 ± 0.9
-0.071
1.1 ± 1.5
0.9 ± 1.3
0.061
1.3 ± 1.7
2.3 ± 4.3
-0.045
4.0 ± 3.2
3.6 ± 2.2
-0.044
1.8 ± 1.6
1.7 ± 1.9
-0.012
42.7 ± 17.5
42.5 ± 22.1
-0.003
1.3 ± 1.3
1.6 ± 2.8
2.6 ± 9.9
0.0 ± 0.0
0.0 ± 0.2
0.0 ± 0.0
0.0 ± 0.0
0.0 ± 0.0
0.0 ± 0.0
0.0 ± 0.0
55
Unit
category (1-4)
%
cm
%
cm2
meters
cm
%
%
%
%
cm
cm
cm
cm
%
%
Scale
meters
%
%
Index
Index
Index
%
%
Index
%
Index
%
count
count
Index
%
cm
count
%
count
cm
count
%
%
cm
%
%
count
Appendix F: Within Meadow DFA λ, r2, and Class Mean Values
Class Means
Meadow
7A
Cave Creek
Flog Salvage
Grapefern
Lost
Midway
Muddy
Peterson Prairie
Smith Butte
λ
80.2
16.2
11.9
96.2
17.3
12.5
554.5
9.7
4.6
2
r
0.98
0.94
0.92
0.99
0.95
0.93
0.99
0.91
0.82
Oviposition
8.8
-4
3.4
9.6
-4.1
3.5
23.2
3.1
-2.3
56
Random
-8.8
3.9
-3.4
-9.6
4.1
-3.5
-23.2
-3.1
2.3
Appendix E: Temperature and Relative Humidity
Data from four iButton data loggers placed in six Mardon skipper Meadows,
Standard deviation = SD. Temperature in degrees Celsius.
Meadow
7A
Season
Fall
Winter
Spring
Summer
Bunny Hill
Fall
Winter
Spring
Summer
Flog
Fall
Winter
Spring
Summer
Grapefern
Fall
Winter
Spring
Summer
Lost
Fall
Winter
Spring
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Temp
RH
Mean
4.04
100.44
0.00
112.01
-0.17
112.25
11.81
92.68
10.39
48.76
0.22
110.71
3.15
106.59
17.18
38.09
6.36
94.08
0.06
110.96
0.77
111.21
0.77
111.21
4.53
98.43
4.53
98.43
-0.16
82.84
-0.16
82.84
5.87
95.29
14.15
82.91
1.07
109.71
57
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
±
SD
6.71
18.70
0.30
1.62
0.27
2.31
10.33
23.73
11.14
43.83
0.27
1.24
7.50
15.41
12.24
34.61
11.39
23.03
0.11
1.73
3.70
6.71
12.74
25.22
7.09
19.12
0.26
39.07
0.26
39.07
10.54
29.98
7.77
21.51
0.24
1.62
3.95
6.70
Minimum Maximum
-5.99
39.63
11.13
113.83
-0.44
0.60
109.42
115.98
-0.44
0.60
107.93
116.46
-1.96
46.59
14.13
115.50
-5.48
50.04
-3.93
112.52
-0.47
0.60
106.82
113.98
-3.42
47.62
23.58
114.98
-0.94
56.03
2.61
114.35
-16.51
44.64
15.50
116.39
-0.43
0.10
106.35
115.38
-5.45
35.67
25.12
118.41
-6.45
41.60
20.78
118.91
-6.41
37.20
12.54
114.08
-0.46
0.13
29.72
112.43
-0.44
0.13
-4.22
112.92
-3.39
43.61
-21.59
111.69
-5.97
36.13
13.58
113.95
-0.44
0.61
105.64
113.88
-0.94
32.15
37.80
114.50
Meadow
Peterson
Season
Summer Temp
RH
Fall
Temp
RH
Winter
Temp
RH
Spring
Temp
RH
Summer Temp
RH
Mean
14.15
82.91
5.60
100.87
0.13
108.51
0.95
105.13
14.62
89.28
58
±
±
±
±
±
±
±
±
±
±
SD
9.52
25.78
9.08
18.92
0.16
7.36
3.38
13.40
11.10
23.27
Minimum Maximum
-1.44
41.62
13.94
111.44
-9.46
47.08
20.22
116.92
0.06
1.08
89.27
116.40
-0.90
29.17
37.56
115.89
-4.93
48.12
15.16
113.29

oviposition selection by a rare grass skipper, polites mardon, in

Transcription

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